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GREAT BASIN NATURAUST MEMOIRS

Brigham Young University

Number 12

MC2
LIBRARY

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Research inthe mrd
Auchenorrhynchai'"''''
Homoptera:
A Tribute to Paul W, Oman

GREAT BASIN NATURALIST

Editor. Stephen L. Wood, Department of Zoology, 290 Life Science Museum, Brigham Young
University, Provo, Utah 84602.

Editorial Board. Kimball T. Harper, Chairman, Botany and Range Science; Perron L. An-
dersen, Zoology; James R. Barnes, Zoology; Hal L. Black, Zoology; Jerran T. Flinders,
Botany and Range Science; Stanley L. Welsh, Botany and Range Science. All are at
Brigham Young University.

Ex Officio Editorial Board Members include Bruce N. Smith, Dean, College of Biological and
Agricultural Sciences; Norman A. Darais, University Editor, University Publications;
Stephen L. Wood, Editor, Great Basin Naturalist.

The Great Basin Naturalist was founded in 1939. The journal is a publication of Brigham
Young University. Previously unpublished manuscripts in English pertaining to the biological
natural history of western North America are accepted. The Great Basin Naturalist Memoirs
series was established in 1976 for scholarly works in biological natural history longer than can be
accommodated in the parent publication. The Memoirs appears irregularly and bears no
geographical restriction in subject matter. Manuscripts for both the Great Basin Naturalist and
the Memoirs series will be accepted for publication only after exposure to peer review and
approval of the editor.

Subscriptions. Annual subscriptions to the Great Basin Naturalist are $25 for private
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$15 for student subscriptions. The price of single issues is $12. All back issues are in print and
are available for sale. All matters pertaining to subscriptions, back issues, or other business
should be directed to the Editor, Great Basin Naturalist, Brigham Young University, 290 Life
Science Museum, Provo, Utah 84602. The Great Basin Naturalist Memoirs may be purchased
from the same office at the rate indicated on the inside of the back cover of either journal.

Scholarly Exchanges. Libraries or other organizations interested in obtaining either journal
through a continuing exchange of scholarly publications should contact the Brigham Young
University Exchange Librarian, Harold B. Lee Library, Provo, Utah 84602.

Manuscripts. See Notice to Contributors on the inside back cover.

10-88 1100 36917

ISSN 017-3614

GREAT BASIN NATURALIST MEMOII

Brigham Young University

dumber 12

Research in the

Auchenorrhyncha,

Homoptera:

A Tribute to Paul W. Oman

CONTENTS

Preface. Mervin W. Nielson 1

Paul W. Oman — an appreciation. John D. Lattin 3

Some new cicadellids from grasslands of Karachi, Pakistan (Homoptera: Cicadellidae).

Manzoor Ahmed, A. Qadeer, and K. F. Malik 10

Classification of the subgenus Athijsanella , genus Athysanella Baker (Homoptera:

Cicadellidae: Deltocephalinae). H. Derrick Blocker and James W. Johnson 18

A new genus, lleopeltus, related to Chlorotettix (Homoptera: Cicadellidae). Paul S.

Cwikla 43

Some aspects of the biology, morphology, and evolution of leaflioppers (Homoptera:

Cicadelloidea and Membracoidea). J. W. Evans 61

Revision of the genus Calliscarta (Homoptera: Cicadellidae: Neobalinae). Paul H.

Freytag 67

A new species oiPazu Oman from eastern California (Homoptera: Cicadellidae: Delto-
cephalinae). Raymond J. Gill 82

Evidence for an Indo-Pacific origin of Hawaiian endemics in Balcltitha and related genera

(Cicadellidae: Macrostelini). W. J. Knight and M. D. Webb 86

Colladonus and related genera of Mexico and Central America with new taxa and

synonymy (Homoptera: Cicadellidae). M. W. Nielson 103

New World Fulgoridae, Part I: genera with elongate head processes. Lois B. O'Brien . . 135
A numerical taxonomic analysis of interspecific morphological differences in two closely

related species of Cicada (Homoptera, Cicadidae) in Portugal. J. A. Quartau 171

Revision of the Nirvaninae (Homoptera: Cicadellidae) of the Indian subcontinent. C. A.

Viraktamath and C. S. Wesley 182

Genus Flexa7nia: new species, phylogeny, and ecology. Robert F. Whitcomb and Andrew

L. Hicks .' 224

Afronisia, a new African genus of Meenoplidae (Homoptera: Fulgoroidea). Michael R.

Wilson 324

Delphacidae of Alaska (Homoptera: Fulgoroidea). Stephen W. Wilson 335

Taxa in the Homoptera, Auchenorrhyncha described by Paul W. Oman 344

Index 349

Great Basin Naturalist Memoirs

Research in the

Auchenorrhyncha,

Homoptera:

A Tribute to

Paul W. Oman

No. 12

Brigham Young University, Provo, Utah

1988

PREFACE

A committee was formed late in 1986 to
organize and develop a volume of taxonomic
papers on the Auchenorrhyncha, Homoptera
to honor Dr. Paul W. Oman for his outstand-
ing contributions in leafhopper systematics.
This volume is the result of efforts by the
committee and by contributors to the work.
The 15 papers form not only a dedicatory trea-
tise of Auchenorrhyncha systematics, but also
a printed emblem of esteemed appreciation to
Dr. Oman for his untiring efforts and many
years of dedicated service in promoting leaf-
hopper systematics in particular and the ento-
mological sciences in general.

The biography of Dr. Oman, prepared by
his long-time associate John Lattin, and a spe-
cial review of the biology, morphology, and
evolution of leaftioppers by the Australian tax-
onomist, John Evans, comprise part of the
volume. The remainder consists of a selected
group of 10 taxonomic papers on leaffioppers,
three on planthoppers, and one on cicadas.
The scope of these papers ranges from the

description of a single taxon to a full taxonomic
treatment of a higher category (subfamily).
Each paper offers important data on the bio-
geography of the respective group. Others
cover host plant associations and relation-
ships, phylogeny, and numerical taxonomy.

Authors of the volume are long-time friends
and associates of Dr. Oman, including a for-
mer graduate student. Dr. C. A. Viraktamath.
A symposium organized by H. Derrick
Blocker and Paul H. Freytag was held at the
1988 Annual Meeting of the Entomological
Society of Louisville, Kentucky. The culmina-
tion of this symposium was the keynote ad-
dress by Robert F. Whitcomb, the presenta-
tion of papers, and the presentation of abound
copy of this volume to the honoree by Mervin
W. Nielson.

The committee expresses appreciation to
Dr. Stephen L. Wood, editor of The Great
Basin Naturalist, Brigham Young University,
for his efforts in editing and preparing the
volume for publication. We also acknowledge

Great Basin Naturalist Memoirs

No. 12

Mrs. Terry Simmons and Mrs. Karen Jensen,
Monte L. Bean Museum, Brigham Young
University, for their invaluable assistance in
typing those manuscripts that required con-
version to the IBM WordPerfect system.

Mervin W. Nielson,
Coordinator

H. Derrick Blocker
Leon W. Hepner
Robert F. Ruppel

Paul H. Freytag
James P. Kramer

Organizing Committee

PAUL W. OMAN— AN APPRECIATION

John D. Lattin'

Abstract — The contributions to professional entomology made by Paul W. Oman are reviewed. A bibliography of
his published contributions to this field from 1930 to 1987 is included.

I first met Paul Oman in December 1950 in
Denver, Colorado, at the national meeting of
the Entomological Society of America. He
was in the uniform of the U.S. Army with the
rank of major, having been called up again to
serve in the Korean War (or "ruckus" as Paul
preferred to call it). I was a graduate student at
the University of Kansas, working with H. B.
Hungerford. Dr. Hungerford encouraged me
to attend the meeting, as did the other faculty
members. He took special care to introduce
the graduate students to other entomologists
at the meeting, including Paul Oman, himself
a graduate of the University of Kansas. My
recollection of that meeting was that Paul took
special interest in each student he met, even
though his time was limited and he was quite
busy with society affairs. He still takes time to
meet and work with students. Now, thirty-
eight years later, we share the same depart-
ment, coffee room. Systematic Entomology
Laboratory (OSU), and a good many hours of
collaboration and discussion in and out of the
field.

Paul Wilson Oman was born in Garnett,
Anderson County, Kansas, on 22 February
1908. He grew up on a farm about five miles
outside of Garnett. His early education was in
a rural school close to his home. He had to
commute the five miles when he later entered
Garnett High School. While growing up, he
made the usual collections of natural history
objects, including insects. A high school biol-
ogy teacher was particularly influential in
keeping alive Paul's interest in the natural
sciences.

He first attended the University of Kansas
at Lawrence to strengthen his background in
mathematics and English, since he was con-
sidering the possibility of a career in the Navy,

including attending Annapolis. He took a
course in entomology to satisfy a biological
science requirement and soon transferred to
that department. Among the departmental
faculty were H. B. Hungerford, chairman,
K. C. Doering, P. B. Lawson, R. H. Beamer,
and P. A. Readio. It is interesting to note that
Hungerford (my own major professor in 1950)
worked on aquatic Hemiptera, Readio on the
Reduviidae, Kathleen Doering was a mor-
phologist but worked on Homoptera, and
both Lawson and Beamer worked not only on
Homoptera but also on leafhoppers. Not sur-
prisingly, Paul's interest in this group of in-
sects was kindled at K.U., and he has contin-
ued to work on the family during his entire
scientific career.

Paul Oman made a fine academic record at
the University of Kansas, being elected to
Sigma Xi, Phi Sigma, and Phi Beta Kappa in
1930. Prior to that date he had been elected to
the Pen and Scroll in 1927 — an honorary soci-
ety in the English Department. Those who
have read Paul's papers or corresponded vdth
him know that this award was well deserved
and know too that he likes to communicate via
the written memo. He graduated from the
University of Kansas with an A.B. in entomol-
ogy in the spring of 1930.

Paul joined the recently organized Taxo-
nomic Unit of the Bureau of Entomology,
U.S. Department of Agriculture, in October
1930. This was to be a long association, for he
retired from the USDA in 1967 when he
joined the faculty of the Department of Ento-
mology at Oregon State University. His re-
sponsibilities included the auchenorrhync-
hous Homoptera and the Psyllidae. Thus, his
interests in the Homoptera, fostered by the
faculty at the University of Kansas, provided

Systematic Entomology Laboratory, Department of Entomology, Oregon State University. Corvallis, Oregon 97331.

Great Basin Naturalist Memoirs

No. 12

Fig. 1. Paul W. Oman.

his entree into his profession. The need for a
speciahst in the "hard" Homoptera was
prompted by the extensive ecological work
being done on the beet leafliopper and associ-
ated insects, especially in the Twin Falls,
Idaho, region. This fortuitous association with
Circulifer tenellus (Baker), a vector of "curly-
top of beets in the western United States,
also provided a basis of his lifelong interest in
applied systematics — the application of sys-
tematic techniques toward the solution of
problems of concern to society. His ultimate
discovery that this insect was native to the
Middle East (rather than South America as
some thought) (Oman 1936, 1948) lead to ex-
plorations for biological control agents from
the Middle East. Further, this discovery
strengthened his conviction that systematics
should play an important role in biological
control. Some years later, when he assumed
charge of the entomological taxonomic unit of
the US DA, he was instrumental in adding the
biological control unit, resulting in the joint

organization we see today.

During his early years in Washington,
D.C., he attended evening classes at George
Washington University where he majored in
entomology. He had extra courses from the
University of Kansas, and these, together
with his course work transferred to Lawrence
from George Washington, plus a thesis, re-
sulted in an M.A. degree in entomology from
the University of Kansas in 1935. Not surpris-
ingly, his thesis was on the leafhoppers and
entitled "A Generic Revision of American
Bythoscopinae and South American Jassinae,"
later published in the well-known University
of Kansas Science Bulletin (Oman 1938). He
continued his academic work at George
Washington University as time permitted and
received a Ph.D. degree from that institution
in 1941. His thesis, somewhat revised, ap-
peared under the title, "The Nearctic Leaf-
hoppers (Homoptera: Cicadellidae), a Gener-
ic Classification and Check List" (Oman 1949).
As Z. P. Metcalf stated (1950, Ann. Entomol.
Soc. Amer. 43: 458), "This is one of the most
outstanding recent contributions to the study
of one of the most difficult families of the
Homoptera, and indeed it was. Paul pro-
duced this landmark publication by working
quietly, steadily, and efficiently, and accom-
plishing a great deal — a Paul Oman hallmark.
He was working as a taxonomic specialist at
this time, and so the courses and thesis work
were done largely after hours. He was the
scientist responsible for research on the
Auchenorrhyncha. The hiatus between the
completion of his dissertation and the appear-
ance of the published version was caused by
World War II.

Paul Oman entered the U.S. Army as a first
lieutenant in the Medical Corps in October
1942 and was assigned to the South Pacific and
the Far East commands. He served in that
area, being promoted to captain, until he re-
turned and left active service in February
1946. While in the South Pacific, he was re-
sponsible for the organization and direction of
entomological activities on Guadalcanal (No-
vember 1943-February 1945) and Okinawa
(April-June 1945). Although other medical
entomology problems were involved, the con-
trol of malaria vectors was the chief effort in
both locales. The details of some of these ac-
tivities can be found in Oman and Christen-
son (1947) and Harper, Downs, Oman, and

1988

Lattin: PaulW. Oman

Levine (1963). At the time of his discharge
from the U.S. Army and his return to the
Division of Insect Identification, US DA, he
could hardly anticipate that the experience
gained in medical entomology would be used
again — in 1950, when he was recalled into the
Army to serve three years in the Korean War.

During the four years following his return
from the South Pacific, Paul was deeply in-
volved in the auchenorrhynchous Homop-
tera. He became project leader in the Hem-
iptera and ultimately assistant division leader
of the Division of Insect Identification. He
published a number of papers, including his
253-page generic classification of the Nearctic
leaflioppers, as well as an account of some of
the medical entomology work done in the
South Pacific. His interest in applied system-
atics is reflected in the papers of this period.

In September 1950 he was reactivated to
regular duty in the U.S. Army at the rank of
major and assigned to the Far East during the
Korean War. His first assignment was as ento-
mologist. Headquarters, 3rd Army. Later he
became chief of the Department of Entomol-
ogy, 406th Medical General Laboratory in
Tokyo, and then commanding officer. Far
East Medical Research Unit, the position he
held until his discharge in August 1953. He
published several papers dealing with medi-
cal entomology during this time, and, of spe-
cial note to homopterists, a paper describing
three new species of Errhornus with a key to
the species. I doubt that he ever thought he
would devote many of his later years to an
intensive study of the genus Errhomus (Oman
1987), but we are glad he did.

Once again he was discharged from the
U.S. Army and again returned to the U.S.
Department of Agriculture, this time as the
head of what became the Insect Identification
and Parasite Introduction Research Branch,
Agricultural Research Service. His systematic
work continued, as did his deep involvement
in all aspects of entomology, entomological
administration, and the Entomological Soci-
ety of America. Gradually, his writing began
to reflect his increased responsibilities and his
ever-broadening interests, but always there
was a deep interest in the leaflioppers.

Part of his administrative responsibilities
involved increased international activities.
He was the leader of the United States ento-
mological delegation to the USSR in 1959 un-

der the U.S. -USSR Scientific and Cultural
Exchange Program. In October 1960 he
moved to New Delhi, India, to become the
director. Far East Regional Research Office,
Foreign Research and Technical Programs
Division, ARS, USDA. There he was respon-
sible for all technical and administrative as-
pects of agricultural research studies in Asiatic
countries where Public Law 480 funds were
available for research of mutual interest to the
United States and the foreign country.

Paul returned home from India in Decem-
ber 1962 to become assistant to the director.
Entomology Research Division, ARS, USDA.
A year later he was appointed assistant direc-
tor of the Entomology Research Division, a
position he retained until he retired from the
U.S. Department of Agriculture in 1967.
Paul's responsibilities included all extramural
research programs, chairman of the Division
Committee for the evaluations of research
personnel, administration of divisional labora-
tories, and an active role in the planning and
development of programs and policies of the
Entomology Research Division. Still he con-
tinued to publish, chiefly on topics related to
his responsibilities with the Division, but he
found time to initiate and participate in the
first conference under the new U.S. -Japan
Cooperative Science Program. This confer-
ence, held in Japan, dealt with arthropod-
borne plant viruses (Maramorosch and Oman
1966). He was responsible for another U.S.-
Japan Conference held in Washington, D.C.,
in 1967. This time the subject matter was
"Systematics in Relation to the Geographical
Distribution of Insects in the Pacific. " Paul
Oman and Karl V. Krombein were the orga-
nizers for the United States. It was obvious
that Paul was held in high regard by his
Japanese colleagues. I was pleased to have
been a participant in that gathering; it was a
special occasion. By this time Paul had retired
after 37 years with the U.S. Department of
Agriculture and had joined the faculty of the
Department of Entomology at Oregon State
University, and yet another chapter had be-
gun in his productive career. He still main-
tains active connections with the USDA, and,
even as this is being written, he is assisting
them to resolve a problem dealing with the
Homoptera.

Paul W. Oman, now Professor Paul W.
Oman, joined the departmental faculty in the

Great Basin Naturalist Memoirs

No. 12

fall of 1967. He assumed many of my responsi-
bilities in systematic entomology when I
moved into administration. Characteristi-
cally, he launched into his new career with
great energy. He developed his own course in
systematic entomology, developed a course in
advances in pest management, and began to
direct graduate student studies. His enor-
mous experience in all aspects of entomology
made him a highly valued member of the
department, and his advice was sought (and
still is) on many topics. As program director at
Oregon State University, he participated in
the NSF-funded Inter-University Program in
Pest Population Ecology that ran from July
1969 until August 1975, a forerunner of the
IPM programs at OSU we know today. Most
of the leading ecologists of the world were
brought to our campus (and other campuses as
well) under this program. It was obvious that
pest problems were universal problems, and
solid science was required to solve them.
During this time educational turmoil was ev-
erywhere, but these well-organized programs
provided some academic stability during un-
stable times. The Pest Population Ecology
program provided a solid core upon which was
built a revised curriculum.

It took him less than a year to obtain fund-
ing from the National Science Foundation for
his work on the systematics of the lealhop-
pers. This funding extended from 1968 until
he retired in 1975. Again, his publications
reflected his efforts and interests as he pub-
lished a series of leaffiopper papers. At the
same time, he was publishing papers dealing
with other aspects of his activities. He also
renewed his interest in the genus Errhomus
and spent many seasons in the field carefully
documenting the complex systematic and bio-
geographical problems associated with that
taxon. The result, of course, is the superb
monograph on Errhomus that appeared in
1987. While many people would be content to
stop there, even before the publication was in
press he took up the world catalog project
with Bill Knight and Merv Nielson. Compla-
cency is not a characteristic of Paul Oman.

Paul was curator of the Entomology Mu-
seum from 1967 to 1971. It must have seemed
like child's play after having been responsible
for the entire systematic operation of the
USDA, and yet he took it very seriously and
made major strides in establishing sound pro-

tocol for the management of the collection.
He added many needed books and sought out
reprints from major systematists. An active
effort was made to acquire, mount, label, and
accession thousands of specimens. He made a
special effort to develop the Homoptera col-
lection that had languished. During the years
since his release as official curator, he has
added literally tens of thousands of specimens
to the collection, and still continues to add
more. These specimens are all mounted, la-
beled, and, if leaflioppers, identified, and cu-
rated. Talk about a curator's dream! We all
know that collections become major scientific
resources because of the work of many people
with a common goal. Paul is the epitome of
such a person.

During the ensuing years he served ento-
mology in many ways. He was on the editorial
board of the Annual Review of Entomology
from 1972 to 1976. He was chairman of the ad
hoc Committee on Entomological Collections
in the United States. He was a member of the
Entomological Society of America Advisory
Committee on Systematic Resources in Ento-
mology from 1973 to 1975. He was a member
of the National Policy Advisory Committee
(NSF) for the National Drosophila Species Re-
source Center in 1975. He was secretary of a
study team that prepared a 138-page report in
1978 entitled "Biological Agents for Pest Con-
trol: Status and Prospects ' for the USDA in
cooperation with land-grant universities, the
State Department of Agriculture, and the
Agricultural Research Institute.

When his long-time friend and colleague,
Knud Swenson, then chairman of our depart-
ment, was forced to step down because of a
tragic illness, Paul took over as acting chair-
man. He served in that capacity from 1973 to
1974 while the department sought a new de-
partment chair. His long experience in sci-
ence administration was most evident during
that time, and we were able to experience yet
another facet of his skills. He was a low-key,
but a most efficient and effective, administra-
tor. He paid attention to detail and provided
leadership and stability that created an atmo-
sphere of professionalism. He rarely com-
plained and favored working out a solution to a
problem. Because his dedication to the job at
hand was always evident, people responded
in a similar fashion. He was especially effec-
tive in getting the most out of the resources
available to him.

1988

Lattin: Paul W. Oman

Paul Oman retired in 1975 — again. There
was only an imperceptible change before and
after this date — occasionally he would head
for the golf course if the afternoon was espe-
cially nice. Although I have never played golf
with Paul, I know some who have — be pre-
pared for a fierce competitor and don t wager
very much on each hole. During the past 13
years of his "retirement" he has averaged six to
eight hours a day in his office, has spent hun-
dreds of hours in the field throughout the
West, has given many reading and conference
courses to many students, has served on nu-
merous graduate student committees, has
hosted foreign scientists and freely shared
with them his vast experience with the Ci-
cadellidae, and has identified thousands of
leafhoppers for many individuals and institu-
tions.

One of my most recent interactions with
Paul occurred on 16 March 1988, when he
participated in the program planning session
for Adam Asquith, one of my new doctoral
students. Paul had given Adam a reading and
conference course in zoological nomenclature
during the preceding three months. In fact,
one of the other graduate committee mem-
bers brought a particularly knotty problem on
nomenclature to the meeting. Ultimately, the
problem was resolved by Paul and Adam. Paul
brought several current articles from science
that were appropriate to the graduate pro-
gram and the proposed thesis topic. He has
always given help freely to the students and
faculty, and he did so again on 16 March 1988.
His eightieth birthday had been celebrated
three weeks earlier, and he had supposedly
retired in 1975, but for Paul Oman it was just a
regular day.

As one who has known him since 1950, 1 can
attest to his many contributions and deep
devotion to entomology. He joined the ESA
in 1929, served as president in 1959, was
elected an honorary member in 1975, and
received the Woodworth Award from the
Pacific Branch in 1982. He joined our faculty
in 1967, and while he holds the title Emeritus
Professor of Entomology, his efforts are indis-
tinguishable from those of regular faculty
members. He is one of those rare scientists
whose influence has extended far beyond his
office and laboratory. I cherish our friendship.

Paul Wilson Oman Bibliography

L Oman, P. W. 1930. A new Paracoelidia (Homoptera:
Cicadellidae). J. Kansas Entoniol. Soc. 3; 78-79.

2. Oman, P. W. 193L Some new Neocoelidia with notes

on other species (Homoptera: CicadelHdae). J.
Kansas Entomol. Soc. 4: 62-68.

3. Oman, P. W. 1931. Newjassinae, with notes on other

species. J. Washington Acad. Sci. 21: 430-436.

4. Oman, P. W. 19.32. Two new Laevicephalus from

CaUfornia. Proc. Entomol. Soc. Washington 34:
90-92.

5. Oman, P. W. 19.33. A classification of North American

agallian leaflioppers. U.S. Dept. Agric. Tech.
Bull. 372. 93 pp.

6. Oman, P. W. 1933. Phlepsius ishidae Matsumura in

North America. Proc. Entomol. Soc. Washington
35: 205-206.

7. Oman, P. W. 19.34. New species and a new genus of

deltocephaline leaflioppers (Hemiptera: Ho-
moptera). Proc. Entomol. Soc. Washington 36:
75-81.

8. Oman, P. W. 1934. South American leafhoppers of

the genus Agalliana. Rev. de Entomol. 4:
333-340.

9. Oman, P. W. 1934. The agallian leafhoppers of the

Biologia material. Ann. Entomol. Soc. Amer. 27:
445-461.

10. Oman, P. W. 1935. New North American agallian

leafhoppers with notes on other species. J. Kansas
Entomol. Soc. 8: 9-16.

11. Oman, P. W. 1936. New Neotropical empoascan

leaflioppers. J. Washington Acad. Sci. 26: 34-40.

12. Oman, P. W. 1936. A new Poblicia from Texas (Ho-

moptera: Fulgoridae). J. Kansas Entomol. Soc. 9:
105-107.

13. Oman, P. W. 1936. Two new leaflioppers from tropi-

cal America. Pan-Pac. Entomol. 12: 116-119.

14. Oman, P. W. 1936. Distributional and synonymical

notes on the beet leafhopper, Eutettix tenellus
(Baker). Proc. Entomol. Soc. Washington 38:
164-165.

15. Oman, P. W. 1936. South American leafhoppers of

the genus Xerophloea Gerniar (Homoptera). Rev.
de Entomol. 6: 394-401.

16. Oman, P. W. 1937. Fossil Hemiptera from the Fox

Hills Sandstone (Cretaceous) of Colorado. J. Pale-
ontol. 11:38.

17. Oman, P. W. 1937. The leaflioppers of the genus

Scaphoidula (Homoptera: Cicadellidae). Rev. de
Entomol. 7: 92-96.

18. Oman, P. W. 1937. The cinerosus group of the genus

Laevicephalus (Homoptera: Cicadellidae). J.
Washington Acad. Sci. 27: 474-479.

19. Oman, P. W. 1937. New eupterygine leafhoppers

from Puerto Rico (Homoptera: Cicadellidae).
Univ. Puerto Rico J. Agric. 21: 567-.572.

20. Oman, P. W. 1938. A generic revision of the Ameri-

can Bythoscopinae and South American Jassinae.
Univ. Kansas Sci. Bufl. 24: 343-420.

21. Oman, P. W. 1938. A contribution to the classification

of South American agallian leafhoppers. Ann.
Carnegie Museum 25: 351-460.

22. Oman, P. W. 1938. Revision of the Nearctic leafliop-

pers of the tribe Errhomenellini (Homoptera:

Great Basin Naturalist Memoirs

No. 12

Cicadellidae). Proc. U.S. National Museum 85:
16.3-180.

23. Oman, P. W., and Nancy H. Wheeler. 19.38. The

North American empoascan leafhoppers of the as-
persa group (Homoptera; Cicadellidae). Proc. En-
tomol. Soc. Washington 40: 1,33-147.

24. Oman, P. W. 1939. Revision of the genus Cpra<aga//ia

Kirkaldy (Homoptera; Cicadellidae). J. Washing-
ton Acad. Sci. 29: 529-.543.

25. Oman, P. W. 1940. Three new species of Delto-

cephalus (Homoptera: Cicadellidae). Proc. Ento-
mol. Soc. Washington 42: 201-203.

26. Oman, P. W. 1941. Revision of the Nearctic Megoph-

thalminae (Homoptera: Cicadellidae). J. Wash-
ington Acad. Sci. 31: 203-210.

27. Oman, P. W. 1943. A new leafliopper from Oceania

(Homoptera: Cicadellidae). Pan-Pac. Entomol.
19: 33-34.

28. Oman, P. W. 1943. A new leafliopper of the genus

Helochara (Homoptera: Cicadellidae). Proc. En-
tomol. Soc. Washington 43: 74-75.

29. Oman, P. W., and R. H. Reamer. 1944. Some new

species of Cuerna (Homoptera: Cicadellidae). J.
Kansas Entomol. Soc. 17: 119-128.
.30. Oman, P. W., and Arthur D. Cushman. 1946. Collec-
tion and preservation of insects. U.S. Dept. Agric.
Misc. Publ. 601. 42 pp.

31. Oman, P. W. 1947. The types of auchenorrhynchous

Homoptera in the Iowa State College collection.
Iowa State College J. Sci. 21: 161-228.

32. Oman, P. W. 1947. Miscellaneous notes on Cicadelli-

dae (Homoptera). J. New York Entomol. Soc. .55:
.59-63.

.33. Rlack, L. M., and P. W. Oman. 1947. Parthenogene-
sis in a leafhopper, Agallia quadripunctata (Pro-
vancher) (Homoptera: Cicadellidae). Proc. Ento-
mol. Soc. Washington 49: 19-20.

.34. Oman, P. W., and L. C. Christenson. 1947. Malaria
and other insect-borne diseases in the South
Pacific campaign, 1942-1945. III. Entomology,
Amer. J. Trop. Med. 27: 91-117.

.35. Oman, P. W. 1948. Notes on the beet leafhopper,
Circiilifer tenellus (Raker), and its relatives (Ho-
moptera: Cicadellidae). J. Kansas Entomol. Soc.
21: 10-14.

36. Oman, P. W. 1948. Distribution of Baldulus maidis
(DeLong and Wolcott) (Homoptera: Cicadellidae).
Proc. Washington Entomol. Soc. .50: 34.

.37. Oman, P. W. 1949. A leafhopper injurious to culti-
vated prune in the western United States. J. Econ.
Entomol. 41:983.

.38. Oman, P. W. 1949. The Nearctic leafhopper (Ho-
moptera: Cicadellidae). A generic classification
and check list. Memoir No. 3, Entomol. Soc.
Washington. 2.53 pp.

.39. Raker, W. L., P. W. Oman, and Thaddeus J. Parr.
19.50. Sucking insects. Pages 105-152 in Craig-
head, Insect enemies of eastern forests. USDA
Misc. Publ. 6.57.

40. Oman, P. W. 1951. Whv the flv? Pages 24-25 in Med.

Rull. No. Ill, Vol. I, Hdcjrs. 3rd Army.

41. Oman, P. W. 1952. How to collect and preserve

insects for study. Pages 65-78 in Insects. U.S.
Department of Agriculture Year Rook.

42. Oman, P. W. 1952. Three new Errhontiis, with a key

to the species (Homoptera: Cicadellidae). Pan-

Pac. Entomol. 23: 13-15.

43. Oman, P. W. 19.55. Review of. J. Linsley Gressit,

Insects of Micronesia, introduction. Rull. Rrook-
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44. Oman, P. W. 19.57. Under separate cover — . Agric.

Chem. 12: 33-34, 93, 95.

45. Oman, P. W. 19.57. The relation of insect taxonomy to

mosquito control. Mosq. News 17: 147-151.

46. Oman, P. W. 19.59. Strength through union. Rull.

Entomol. Soc. Amer. 5: 1-2.

47. Oman, P. W. 1960. The relation of insect taxonomy to

applied biology. Rull. Entomol. Soc. Amer. 6:
3-5.

48. Oman, P. W. 1961. What insects are positively photo-

sensitive? (Summary). Pages 33-34 in Response of
insects to induced light. ARS 20-10, July.

49. Oman, P. W., Lev. F. Curl, Theodore R. Davich,

Herbert L. Haller, Halbert M. Harris, Carl R.
Huffaker, and John V. Osmun. 1961. Entomology
in the Soviet Lhiion, report of a technical study
group. USDA, ARS. 24 pp.

50. Harper, P. A., W. G. Downs, P. W. Oman, and N. D.

Levine. 1963. Chapter VIII, New Hebrides,
Solomon Islands, Saint Matthias Group, and
Ryukyu Island. Pages 399-495 in Preventive
medicine in World War II. Vol. VI, Communica-
ble diseases. Office of the Surgeon General, De-
partment of the Army, Washington, D.C.

51. Oman, P. 1964. World problems in entomology in

"Farmer's World." Pages .546-5.54 in U.S. Depart-
ment of Agriculture Year Rook.

.52. Maramorosch, K., and P. Oman. 1966. U.S. -Japan
joint conference on arthropod-borne plant virus-
es. Riosci. 16: 608-610.

.53. Oman, P. 1966. Current trends and broad objectives
of USDA Entomology Research Division Pro-
gram. Proc. North Central Rranch, Entomol. Soc.
Amer. 21: 13-19.

54. Oman P. 1966. Facts and concepts of insect control

without pesticides. Pages 52-62 in C. W. Mc-
Conib, Minutes of the Eastern Plant Roard 4Ist
Annual Meeting. Virginia Reach, Virginia, 26-30
April 1966.

55. Oman, P. 1968. Prevention, surveillance and man-

agement of invading pest insects. Rull. Entomol.
Soc. Amer. 14: 98-102.

56. Oman, P., and K. V. Krombein. 1968. Systematic

entomology: distribution of insects in the Pacific.
Science 161: 78-79.

57. Oman, P. 1969 Criteria of specificit\' in virus-vector

relationships. Pages 1-22 in K. Maramorosch,
ed.. Viruses, vectors and vegetation. Interscience
Publ., John Wiley, xii 4- 666 pp.

58. Oman, P. 1969. Review of R. G. Fennah, Fulgoroidea

(Homoptera) from New Caledonia and the Lo\alty
Islands. Rull. Entomol. Soc. Amer. 15: 2.59-260.'
.59. Oman, P. 1970. Taxonomy and nomenclature of the
beet leiifliopper, Circulifer tenellus (Homoptera:
Cicadellidae). Ann. Entomol. Soc. Amer. 63:507-
512.

60. Oman, P. 1970. Leaflioppers of the Agalliopsis no-

vella complex (Homoptera: Cicadellidae). Proc.
Entomol. Soc. Washington 72: 1-29.

61. Oman, P. 1970. Leafhoppers of the A^a//(opsi.s' uari-

(d>ilis group with description of a new species
(Homoptera: C^icadellidae). Proc. Entomol. Soc.

1988

Lattin: Paul W. Oman

Washington 72; 30-32.

62. Oman, P. 1970. The Oregon State University insect

collection. Bull. Entomol. Soc. of Amer. 16: 165.

63. Oman, P. 1971. A new Agallia from the western

United States. J. Kansas Entomol. Soc. 44: 325-
372.

64. Oman, P. 1971. The iema\e oiThatuna gilletti Oman,

with biological notes. Proc. Entomol. Soc.
Washington 73: 368-372.

65. Oman, P. 1972. Documentation of biological control

operations. International Organization for Biologi-
cal Control, Western Hemisphere Region Sec.
Newsletters: 1-2.

66. Oman, P. 1972. A new megophthalmine leafhopper

from Oregon, with notes on its biology and behav-
ior (Homoptera: Cicadellidae). Royal Entomol.
Soc. London, J. Entomol. (B)41: 69-76.

67. Oman, P. 1972. The lealliopper subfamily Koebeli-

inae (Homoptera: Cicadellidae). Published as part
of a Festschrift honoring Professor K. Yasumatsu
of Japan, pp. 129-139 (dated 1971, issued 1972).

68. Oman, P. 1974. Curtis P. Clausen ('32). (A vignette.)

Cosmos Club Bull. 27: 3-5.

69. Oman, P. 1974. Identification and classification in

pest management and control. Pages 77-86 in F.
G. Maxwell and F. A. Harris, eds., Proc. Summer
Institute on Biological Control of Plant Insects and
Diseases. University of Mississippi Press, Jack-
son. 647 pp.

70. Oman, P., and C. A. Musgrave. 1975. The Nearctic

genera of Errhomenini (Homoptera: Cicadelli-
dae). Melanderia21: 1-14.

71. Oman, P. 1976. Review of World catalogues of the

Homoptera: Auchenorrhyncha. Bull. Entomol.
Soc. Amer. 22: 161-164.

72. Gill, R. J., and Paul Oman. 1982. A new species and

new distributional records for megophthalmine
leafhoppers, genus Tiaja (Homoptera: Cicadelli-
dae). Entomography 1; 281-288.

73. Klein, M., B. Raccah, and P. W. Oman. 1982. The

occurrence of a member of the Circulifer tenellus
species complex (Homoptera: Cicadellidae: Eu-
scelini) in Israel. Phytoparasitica 10(4): 237-240.

74. Converse, R. H., R. G. Clarke, P. W. Oman, Sr., and

G. M. Milbrath. 1982. Witches' broom disease of
black raspberrv in Oregon. Plant Disease 66(10):
949-951.

75. Lattin, J. D., and P. Oman. 1983. Where are the

exotic insect threats? Pages 93-137 in Charles
L. Wilson and Charles L. Graham, eds.. Exotic
plant pests and North American agriculture. Aca-
demic Press. 498 pp.

76. Oman, P. 1985. A synopsis of the Nearctic

Dorycephalinae (Homoptera: Cicadellidae). J.
Kansas Entomol. Soc. 58: 314-336.

77. Oman, P., and R. 1. Sailer. 1986. The role of catalog-

ing in the advancement of systematics and bio-
geography. Tymbal (Auchenorrhyncha Newslet-
ter) 7: 16-21.

78. Oman, P. 1987. Alary polymorphism in the Cicadelli-

dae and its ecological implications. Pages 55-63 in
M. R. Wilson and L. R. Nault, eds.. Proceedings
of the 2nd International Workshop on Leafhop-
pers and Planthoppers of Economic Importance.
C.A.B. International Institute of Entomology. 368
pp.

79. Oman, P. 1987. The leafhopper genus Errhomus

(Homoptera: Cicadellidae: Cicadellinae): system-
atics and biogeography. Oregon State University,
Department of Entomology, Systematic Entomol-
ogy Laboratory Misc. Pub. No. 1: 1-72.

SOME NEW CICADELLIDS FROM GRASSLANDS OF KARACHI, PAKISTAN
(HOMOPTERA: CICADELLIDAE)

Manzoor Ahmed', A. Qadeer', and K. F. Malik'

Abstract. — A brief survey of grassland lealhoppers of Karachi during 1985 revealed the existence of five new species
of the family Cicadellidae. These include Exitianus major, E. minor. Chiasmus karachiensis , C. lobosa, and
Goniagnathiis bifurcatus .

Recent studies of grassland leafhoppers of
Pakistan by Ahmed (1986), Ahmed and Rao
(1986), and Ahmed and Yunus (1986) showed
that leafhoppers form a very important com-
ponent of the phytophagous fauna of our
grasslands. The leafhopper fauna described
by these and other workers, e.g.. Blocker et
al. (1972) from USA and Theron (1982) from
South Africa, appears to be widely diversified.
Andrezejewska (1984) believed that species
composition of Auchenorrhyncha in grass-
lands depends to a high degree on immigrant
species, which indicates that most of the spe-
cies encountered in grasslands move in from
the neighboring crops when the latter are har-
vested. Thus, grasslands serve not only as
alternate feeding grounds for leafhopper pests
of a number of crops but also as breeding
places for many species of leafhoppers during
their off-crop seasons. Studies of grassland
leafhoppers could therefore be useful from
many aspects.

The leafhoppers described herein have
been studied following the methodology out-
lined by Ahmed (1985).

Exitianus Ball

The genus Exitianus Ball is known from all
major regions of the world. Oman (1949) re-
described the genus and listed the Nearctic
species. Metcalf (1967) listed 35 world spe-
cies, and Ahmed and Rao (1986) recorded £ .
capicola (Kb.) and described E . peshawaren-
sis from Pakistan. During the present study
two more new species of the genus were col-
lected from Karachi, Pakistan, and are de-
scribed below.

Exitianus m^jor Ahmed & Qadeer, n. sp.

(Figs. 1 A-K)

Length. — Male 4.2 mm, female 4.8 mm;
head broad, rounded at anterior apex; median
length of crown nearly equal to or slightly less
than the interocular width; transocular width
of head slightly more than midlength of
crown; ocelli close to mesal margin of eyes on
anterolateral margin; crown with dim brown
markings, transverse band of dark brown
color in middle, rest of crown pale browTi;
frontoclypeus narrowing anteriorly; ante-
clypeus longer than broad, narrowing anteri-
orly; lora broad; pronotum possessing tiny
spots of brown color on disc; scutellum dim
brown, with two brownish spots anterolater-
ally.

Forewings with apex smoothly rounded;
maximum width slightly prior to midlength;
appendix broad, extending around wing apex
up to 4th apical cell; apical cells 5; subapical
cells 3; veins in the basal region possessing
brownish markings all along; hind wings with
4 closed apical cells.

Male. — Plate in ventral view broad in basal
half, narrowed in apical half, possessing 6-7
macrosetae on ventrolateral surface; pygofer
with posterior margin extended into narrow
posterodorsal lobe, with one long and one
small blackish macrosetae on lobe, style in
dorsal view massive, sclerotized in marginal
areas, membranous in middle, with broad,
bluntly rounded lobe, apical extension
spinose, curved laterad; connective Y-shaped
with arms quite close to each other; aedeagus
with preatrium reduced, dorsal apodeme well

Department of Zoology, University of Karachi, Karachi-32, Pakistan.

1988

Ahmed et al. : New Grassland Cicadellidae from Pakistan

FiPs 1 A K Exitianus maior n sp.: A. head and thorax, dorsal view; B, head, ventral view; C, forewing; D, hind
winfE gentf clpLTe S view' F, male plate, ventral view; G, style, dorsal v.ew; H, connective, dorsal v,ew;
I, aedeagus, lateral view; J, aedeagus, dorsal view; K, female seventh sternum.

developed, shaft in lateral view broad in basal
2/3, narrowed apically, smoothly curved dor-
sally at apex.

Female. — Seventh sternum narrowed pos-
teriorly, posteromedian margin sinuate.

Exitianus major appears quite close to E .

capicola (Stal) in venation and shape of aedea-
gus, but differs in its smaller size, shape ot
connective, chaetotaxy of male plate, and to
some extent in the shape of female seventh

sternum.

Holotype (male).— Pakistan: Karachi, grass,

Great Basin Naturalist Memoirs

No. 12

Fig. 2 A-K. Exitianus mmor, n. sp.: A, head and thorax, dorsal view; B, head, ventral view; C, forewing; D, hind
wing; E, genital capsule, lateral view; F", male plates, ventral view; G, style, dorsal view; H, connective, dorsal view;
I, aedeagus, lateral view; J, aedeagus, dorsal view; K, female seventh sternum.

26. VI. 85, Qadeer. Seven paratypes, .same head projected in front, broadly rounded at

data as holotype, all in Zoological Museum,
University of Karachi, Karachi, Pakistan.

Exitianus minor Ahmed & Qadeer, n. sp.

(Figs. 2 A-K)

Length. — Male 4.00 mm, female 4.5 mm;

anterior apex; median length of crown less
than interocular width; transocular width of
head nearly ecjual to transverse width of
pronotum; coronal suture present up to more
than midlength; ocelli on anterolateral margin

1988

Ahmed ETAL.: New Grassland Cicadellidae from Pakistan

of crown, close to eyes; crown with brownish
markings on disc and transverse brownish
band at the level of anterior margin of eyes;
ground color of crown more or less pale
brown; frontoclypeus nearly parallel sided,
narrowed at apex; anteclypeus narrowed ante-
riorly, much longer than broad; lora broad;
pronotum with some tiny spots in middle in
anterior region, with posterior margin nearly
transverse; scutellum pale brown, with oval-
shaped, brownish spots anterolaterally.

Forewing with apex broadly rounded; api-
cal part slightly broader than basal part; ap-
pendix broad, extending up to 5th apical cell;
apical cells 5; subapical cells 3; hind wing with
4 closed apical cells.

Male. — Plate in ventral view, broad
basally, narrowed apically, possessing 4-5
macrosetae near apex on lateral margin, long
row of 9-10 macrosetae from near base to
apex on ventral surface; pygofer with poste-
rior margin directed into posterodorsal broad
and rounded lobe, lobe possessing 4-5
brownish macrosetae; style in dorsal view
massive, membranous in middle, sclerotized
along margins, preapical lobe flattened, al-
most truncate mesally, possessing 3-4 macro-
setae, apical extension prominent, spinose,
extreme apex directed caudolaterad; connec-
tive Y-shaped, with arms smaller than the
stem, converging at their apex; aedeagus in
lateral view, with preatrium reduced, dorsal
apodeme present, appearing broad in dorsal
view, shaft tubular, curved smoothly dorsad,
apical part narrowed.

Female. — Seventh sternum with posterior
margin possessing three lobes, lateral lobes
more projected than the median.

Exitianus fninor appears close to E. ka-
rachiensis Ahmed and E. major described
hitherto in its venation and general pattern of
male genitalia, but differs in its shape of con-
nective and female seventh sternum.

Holotype (male). — Pakistan: Karachi, grass,
12. V. 1985, Qadeer. Thirty paratypes, all in
Zoological Museum, University of Karachi,
Karachi, Pakistan.

Chiasmus Mulsant and Rey

Distant (1908) recorded Chiasmus uzelli
Melichar from Ceylon. Pruthi (1930) de-
scribed three new species from various parts
of India and reassigned Kartwa mustelina
Distant to the genus Chiasmus . The present

description of two new species in Chiasmus is
the first record of the genus from Pakistan.

Chiasmus karachiensis Ahmed &
Qadeer, n. sp.

(Figs.3A-J)

Length. — Male 2.3 mm; head strongly
projected in front, apex of crown narrowed
and rounded; median length of crown much
less than interocular width; transocular width
of head slightly more than transverse width of
pronotum; coronal suture present up to more
than midlength of crown; ocelli on anterolat-
eral margin of crown, close to eyes; color of
eyes blackish; crown possessing blackish
brown dots throughout; ground color of crown
pale yellow; pronotum yellowish, its posterior
margin nearly straight; scutellum dim yellow;
frontoclypeus narrowing anteriorly, blackish
in color throughout, except median, longitu-
dinal stripe of yellow color; anteclypeus black
in middle; lora broad, expanded.

Forewing with apex smoothly rounded;
maximum width of wing slightly prior to mid-
length; appendix broad, extending to 3rd api-
cal cell; apical cells 4; subapical cells 2; hind
wing with 4 closed apical cells.

Male. — Plate in ventral view broad at base,
middle and apical parts much narrowed,
macrosetae 2-3 on ventral surface in apical
half, apex of plate rounded and directed cau-
dad; pygofer in lateral view with posterior
margin narrowed into angular, posterodorsal,
much-projected lobe, processes and setae
usually absent; style in dorsal view membra-
nous in middle, sclerotized along margins,
preapical lobe short, narrow, and angular,
apical extension dark brown in color, spinose
and curved laterad; connective Y-shaped,
with arms longer than the stem, arms apposed
most of their anterior part, forming nearly
rounded space in middle; aedeagus in lateral
view with preatrium reduced, dorsal apo-
deme absent, shaft tubular, somewhat curved
dorsad towards apex, processes absent.

C . karachiensis appears close to C . alata
Pruthi in general appearance as well as pat-
tern of venation and genitalia, but differs in
the shape of pygofer.

Holotype (male). — Pakistan: Karachi, grass,
1.XI.85, Qadeer. One paratype, same data as
holotype, both in the Zoological Museum,
University of Karachi, Karachi, Pakistan.

Great Basin Naturalist Memoirs

No. 12

Figs. 3 A-J. Chiasmus karachiensis , n. sp.: A, head and thorax, dorsal view; B, head, ventral view; C, forewing; D,
hind wing; E, genital capsule, lateral view; F, male plates, ventral view; G, style, dorsal view; H, connective, dorsal
view; I, aedeagus, lateral view; J, aedeagus, dorsal view.

Chiasmus lohata Ahmed & Qadeer, n. sp.

(Figs. 4 A-J)

Length, — Male 2.5 mm, female 3.0 mm;
head projected and narrowed in front, apex
rounded; median length of crown nearly equal
to or slightly less than interocular width;
transocular width of head equal to transverse
width of pronotum; coronal suture present up
to more than midlength; ocelli present on

anterolateral margin of crown, close to eyes;
crown possessing few, dim, brownish spots in
middle along posterior margin; frontoclypeus
broad, narrowed in apical 1/3; anteclypeus
longer than broad; lora broad; pronotum pos-
sessing rows of brownish markings.

Forewing with apical margin smoothly
rounded; apical cells 4; appendix broad,
reaching 3rd apical cell; subapical cells 2;

1988

Ahmed et al.: New Grassland Cicadellidae from Pakistan

Figs. 4 A-J. Chiasmus lobata, n. sp.: A, head and thorax, dorsal view; B, head, ventral view; C, forewing; D, hind
wing; E, genital capsule, lateral view; F, male plate, ventral view; G, style, dorsal view; H, connective, dorsal view;
I, aedeagus, lateral view; J, aedeagus, dorsal view.

veins with brownish shades along their
length; hind wing with 3 closed apical cells.

Male. — Plate in lateral view broad at base,
narrowed in apical half, apex rounded, pos-
sessing row of 2-3 macrosetae on ventral sur-
face in apical half, another row of 7-8 micro-
setae on lateral margin from near base to mid-
length, length of plate more than the pygofer,
apposed mesally at base; pygofer in lateral
view with posterior margin broad, dented,
more projected posterodorsally, disc with
dorsoventral row of 4 macrosetae; anal tube
moderately developed; style in dorsal view

with preapical lobe narrowed, angular, apical
extension thin, spinose, directed laterad;
aedeagus in lateral view with preatrium re-
duced, dorsal apodeme present, shaft tubu-
lar, somewhat curved dorsad in middle, apex
rounded and thin, directed caudad; connec-
tive Y-shaped, with arms much longer than
stem, arms confluent again forming space in
middle.

Chiasmus lobata appears quite close to C .
karachiensis as well as to C . niger Pruthi and
C . jagdishi Pruthi in the pattern of male geni-
talia, but differs from them in the peculiar

Great Basin Naturalist Memoirs

No. 12

shape of pygofer, particularly the posterodor-
sal margin.

HoLOTYPE (male). — Pakistan: Karachi,
15. XII. 85, grass, Qadeer. One paratype,
same data as holotype, both in Zoological
Museum, University of Karachi, Karachi,
Pakistan.

Goniagnathus Fieber

The genus Goniagnathus Fieber was re-
described by Dlabola (1954). Distant (1918)
described three species of the genus from In-
dia. Ahmed and Yunus (1986) reported G.
guttulinervis Kirsch. from Pakistan. Another
species of the genus is described here from
grass.

Goniagnathus bifurcattis Ahmed &

Qadeer, n. sp.

(Figs. .5 A-I)

Length. — Male 5.0 mm; head with ante-
rior margin broadly convex, smoothly
rounded in front; median length of crown
much less than the interocular width; transoc-
ular width of head slightly more than the
transverse width of pronotum; coronal suture
up to more than midlength; ocelli on antero-
lateral margin of head, close to apex; ante-
clypeus longer than broad; lora broad; prono-
tum with few brownish black dots along
anterior margin and on posterior margin;
scutellum grayish black throughout.

Forewing with apex broad and rounded;
maximum width of wing in basal half; ap-
pendix broad, extending to 4th apical cell;
apical cells 5; subapical cells 3; black markings
present in appendix, 2nd, 3rd, and 4th apical
cells, in some discal subapical cells and along
all the veins; hind wing with 4 closed apical
cells.

Male. — Genital segment strongly sclero-
tized, dark brown in color; plates in ventral
view fused completely to form a single ovid
structure, width of plate decreasing smoothly
to rounded apex, with few blackish macrose-
tae and marginal microsetae in apical part;
pygofer in lateral view with posterior margin
broad, nearly truncate, slightly sinuate pos-
terodorsally, with at least three prominent
groups of stout macrosetae, one closely placed
group posterodorsally, one mixed group of
short and long in middle of posterior margin in
middle, and one group in posteroventral re-
gion of disc, microsetae dispersed throughout

pygofer; anal tube moderately long, possess-
ing fine microsetae; posterior margin of
pygofer finely dentate; style in dorsal view
with preapical lobe minutely developed, api-
cal part expanded, with extreme apex
obliquely truncate; connective in dorsal view
rod-shaped, arms indistinct, connective
curved in middle; aedeagus in lateral view
with preatrium reduced, dorsal apodeme well
developed, closely associated with connec-
tive; shaft of aedeagus short, tapered apically,
long processes arising from atrium, subparal-
lel to shaft, exceeding shaft in length; gono-
pore subterminal on dorsal surface.

Goniagnathus bifurcatiis is close to G. gut-
tulinervis Kirschbaum in the general pattern
of male genitalia, particularly the male plate
and aedeagus, but differs in the shape of style,
pygofer, and connective.

HoLOTi'PE (male). — Pakistan: Karachi, grass,
10. XI. 85, Qadeer. One paratype, same data
as holotype, both in Zoological Museum, Uni-
versity of Karachi, Karachi, Pakistan.

Literature Cited

Ahmed. M 1985. Typhlocybinae of Pakistan. Pakistan
Agric. Res. Council, Islamabad. 270 pp.

1986. Some investigations of leafhoppers of grass-
lands and allied crops in Pakistan (in press).

1986. Some investigations of lealhoppers of grass-
lands in Lower Sind, Pakistan (in press).

Ahmed. M , andS Rao. 1986. Some commonly found leaf
and planthoppers on vegetable plants in the sub-
urbs of Peshawar. NWFP, Pakistan (in press).

Andrezejewska. L. 1984. Ecological structure of
Auchenorrhyncha meadow communities under
increasing cultivation. Mitt. Sch. Entomol. Ges.
57(4): 40,5-406.

Blocker, H D . T L Harvey, and J L Launchbaugh.
1972. Grassland leafhoppers. 1. Leafhopper popu-
lations of upland seeded pastures in Kansas. Ann.
Entomol. Soc. Amer. 6.5(1); 166-172.

Distant. \V L 1908. The fauna of British India including
Ceylon and Burma. Rh\nchota4: 1.56-419.

1918. The fauna of British India including Ceylon

and Burma. Rhynchota 7; 1-109.

Dlabola, J 1954. Fauna CSR Svazek-Homoptera. Prace
Ceskoslocenske Akad. 1954: 7-339.

Metcalf, Z. P 1967. General catalogue of the Ho-
moptera. Fasc. VI(IO). 2695pp.

C)nl\n, P W 1949. The Nearctic le;iflioppers (Homop-
tera: Cicadellidae). A generic classification and
check list. Mem. Entomol. Soc. Washington 3:
1-253.

Pruthl H. S 1930. Studies on Indian Jassidae (Ho-
moptera). Mem. Indian Mns. 11(1): 1-68.

Theron. J G 1982. Grassland le;iflioppers (Hemiptera:
Cicadellidae) from Natal, South Africa, with de-
scriptions of new genera and species. Phytophy-
lactica 14; 17-.30.

1988

Ahmed et al.; New Grassland Cicadellidae from Pakistan

Figs. 5 A-I. Goniagnathus bifurcatus , n. sp. : A, head and thorax, dorsal view; B, head, ventral view; C, forewing; D,
hind wing; E, genital capsule, lateral view; H, aedeagus and connective, lateral view; I, aedeagus, dorsal view.

CLASSIFICATION OF THE SUBGENUS ATHYSANELLA, GENUS
ATHYSANELLA BAKER (HOMOPTERA, CICADELLIDAE, DELTOCEPHALINAE)^

H. Derrick Blocker^ and James W. Johnson^

Abstract. — The subgenus Athysanella Baker is reviewed and a possible phylogeny of the group is discussed. A total
of 35 species is recognized. Descriptions, illustrations, and a key to males are presented. Twelve new species are
described: aphoda, cursa, deserta,furnaca, hemijona, krameri, pastora, strobila, stylosa, vativala, and whitcombi are
from the western United States, and marthae is from Mexico.

The subgenera of Athysanella Baker have
been reviewed by Blocker and Johnson
(1988). The nominate subgenus revised
herein contains 35 species, of which 12 are
new. Distribution of this subgenus is primar-
ily in the western half of the United States,
with 4 species recorded from Mexico and 4
from western Canada. The hosts are primarily
shortgrasses.

Blocker (1984) has described the morpho-
logical variation that occurs in both sexes be-
cause of incompletely developed genital
structures. Undeveloped and partially devel-
oped specimens are often difficult to identify.
We have attempted to illustrate sufficient
characters to allow identification of males, but
the user must attempt to make identifications
from a series of specimens that appear to be
fully developed. As an example, the hind tib-
ial spur of partially developed males may be
absent or reduced; the genitalia also may be
reduced in size.

Females are particularly difficult to deter-
mine. It is best to try to associate them with a
male and then check the shape of the posterior
margin of sternum VII. There is much in-
fraspecific variation in specimens of both
sexes. Ball and Beamer (1940) should be con-
sulted for additional and original descriptions;
Blocker and Wesley (1985) and Wesley and
Blocker (1985) should be consulted for distri-
bution in Canada- Alaska and Mexico, respec-
tively.

Depositories

Persons and institutions (acronyms are used
in the text) furnishing material for this study
include (the authors regret any omissions):

UAz - University of Arizona, F. Werner.

CAS - California Academy of Science, P. Arnaud and

N. Penny.
CDA - California Department of Agriculture, R. Gill

and J. Sorensen.
CNC - Canadian National Collection, K. G. A. Hamil-
ton.
IPL - Insect Pathology Laboratory, US DA, Belts-

ville, A. L. Hicks and R. F. Whitcomb.
KSU - Kansas State University.
KU - Snow Entomological Museum, University of

Kansas, R. W. Brooks.
MWN - Collection of M. W. Nielson.
OhSU - Ohio State University, P. S. Cwikla.
OkSU - Oklahoma State University, R. A. Drew.
OrSU - Oregon State University, P. W. Oman.
UBC - University of British Columbia, G. Scudder and

S. Cannings.
USNM - U.S. National Museum, Natural History, J. P.

Kramer.

Athysanella (Athysanella) Baker

Athysanella Baker, 1898; 185. Type species: Athysanella
magdalena Baker, 1898, by original designation.

This subgenus is characterized by the pres-
ence of a hind tibial spur and the absence of a
pygofer process (Blocker and Johnson 1988).
It is a sister of the subgenus Gladionura.

Species of the subgenus Athysanella:

aphoda Blocker, n. sp. Nevada.

aspera Ball and Beamer, 1940. California.

bifida Ball and Beamer, 1940. Colorado, Montana, New

Mexico, North Dakota, South Dakota, Texas,

Wyoming, Canada.
cursa Blocker, n. sp. Arizona, Colorado, New Mexico,

Utah.
deserta Blocker, n. sp. California.
foeda Ball and Beamer, 1940. Wyoming.
fredonia Ball and Beamer, 1940. Arizona, Colorado, New

Mexico, Nevada, Utah.
furnaca Blocker, n. sp. California.
gardenia Osborn, 1930. Colorado, Wyoming.

'Contribution 88-.312-] from the Kansas Agriculture Experiment Station.
^Department of EntomoloKy, Kansa.s State University. Manhattan, Kansas 66506
Department of Entomology, University of Missouri, Columbia, Missouri 65221.

1988

Blocker, Johnson: Subgenus Athysanella

frigida Oshorn, 1930.

glohosa Ball and Beanier, 1940. Arizona.

hemijona Blocker, n. sp. California.

incerta Ball and Beamer, 1940. Colorado, Kansa.s, Mon-
tana, Nebraska, New Mexico, Wyoming.

incongrua Baker, 1898. Colorado, Iowa, Kan.sas, Ne-
braska, New Hampshire, New Mexico, Okla-
homa, South Dakota, Texas, Wyoming.

kadokana Knull, 1951. Colorado, Montana, Nebraska,
South Dakota, Wyoming, Canada.

kanahana Ball and Beamer, 1940. Arizona, Kansas, New
Mexico, Utah.

krameri Blocker, n. sp. California.

laeta Ball and Beamer, 1940. Arizona, New Mexico.

magdalena Baker, 1898. Arizona, California, Colorado,
Montana, New Mexico, Nevada, Oklahoma,
Texas, Wyoming, Mexico.

inarthae Blocker, n. sp. Mexico.

parca Ball and Beamer, 1940. Colorado, New Mexico.

pastora Blocker, n. sp. New Mexico.

plana Ball and Beamer, 1940. California.

planata Ball and Beamer, 1940. California.

robusta Baker, 1898. Colorado, Montana, Nebraska,
North Dakota, Wyoming, Canada.

montana Osborn, 1930.

rostrata Ball and Beamer, 1940. California.

salsa Ball and Beamer, 1940. Kansas, Texas, Mexico.

strobila Blocker, n. sp. California, Idaho, Nevada, Ore-
gon.

stylosa Blocker, n. sp. New Mexico.

supina Ball and Beamer, 1940. Texas.

tenera Ball and Beamer, 1940. Nevada.

(ereforans (Gillette and Baker), 1895. Colorado, Montana,
North Dakota, Nebraska, Utah, Wyoming, Can-
ada.

utahna Osborn, 1930. Arizona, California, Colorado,
Idaho, Nevada, Utah, Oregon, Washington,
Canada, Mexico.

vativala Blocker, n. sp. Nebraska, North Dakota.

whitcombi Blocker, n. sp. Nevada.

yumana Osborn, 1930. Arizona, California.

Key to Athysanella Males, sensu stricto

1. Style rounded apically; pygofer variable;

plates embrowned apically 2

— Style widened, usually bifid apically; pygofer
usually broadly rounded apically; plates vari-
able . ' ' 5

2(1). Pygofer extended apically; connective, plates,

and vertex variable 3

— Pygofer truncate apically; connective shorter
than style; plates truncate apically; vertex
usually with fuscous spots .... gardenia Osborn

3(2). Aedeagus with basal process, unique; plates
truncate apically; connective as long as style;
vertex without fuscous spots .... robusta Baker

— Aedeagus simple or with ventral serrations;
connective shorter than style; vertex usually
with fuscous spots 4

4(3). Aedeagus with shaft simple, elongate; plates

rounded apically, embrowned . . pastora, n. sp.

— Aedeagus with shaft shortened, expanded and
serrate ventrally; plates truncate apically . . .
globosa Ball & Beamer

5(1). Aedeagus with conspicuous apical hook; style
exceeding apex of plate; vertex with or with-
out fuscous spots 6

— Aedeagus with poorly developed or no apical
hook (apical serrated flanges may be present);
style variable; vertex with or without fuscous
spots 22

6(5). Aedeagus with single apical hook 7

— Aedeagus with semicircular apical hook .... 20

7(6). Aedeagus not widest basally (may have crests
at midlength), not tapered apically; plates
rounded apically; vertex with or without fus-
cous spots 14

— Aedeagus widest in basal 1/2, shaft tapered in
apical 1/2, apical hook conspicuous; plates
usually truncate apically; vertex with or with-
out fuscous spots 8

8(7). Aedeagus with distinct lateral tubercles at
base of shaft; style with ventral apical arm
widened, dorsal arm slender; vertex without
fuscous spots planata Ball & Beamer

— Aedeagus without distinct tubercles; style
with ventral and dorsal arms variable; vertex
with or without spots 9

9(8). Style without apical lateral flange, dorsal arm

short, wide; plates rounded apically

krameri, n. sp.

— Style with distinct apical lateral flange or ven-
tral arm expanded laterally, dorsal arm vari-
able, elongate; plates truncate apically 10

10(9). Aedeagus with apex of dorsal apodeme ex-
panded; style with dorsal arm elongate 11

— Aedeagus with dorsal apodeme rounded;
style variable 12

11(10). Style with dorsal arm slender; vertex with no

fuscous spots; smaller in overall size

whitcombi, n. sp.

— Style with dorsal arm more robust; vertex
usually with fuscous spots; larger in size ....
plana Ball & Beamer

12(10). Aedeagus with shaft robust, short; style with

dorsal apical arm longer, variable apically . . 13

— Aedeagus with shaft less robust; style with

dorsal arm rounded apically

aspera Ball & Beamer

13(12). Style with dorsal arm bulbous apically; aedea-
gus with apical hook conspicuous, with slight
processes at base aphoda, n. sp.

— Style with dorsal arm acute apically; aedeagus
with apical hook less developed, no process at
base strobila, n. sp.

14(7). Style bifid apically; aedeagus with apical hook

short; vertex with or without fuscous spots . . 15

— Styles not bifid apically or dorsal arm elon-
gate; aedeagus with apical hook variable; ver-
tex with or without fuscous spots 17

15(14). Aedeagus short, shaft serrate ventrally,
widest at midlength; style without distinct
lateral apical flange; vertex without fuscous
spots supina Ball & Beamer

Great Basin Naturalist Memoirs

No. 12

— Aedeagus longer, shaft not serrate or
widened; style variable; vertex with or with-
out fuscous spots 16

16(15). Style with conspicuous apical lateral flange;

vertex without fuscous spots

foeda Ball & Beamer

— Style without apical lateral flange; vertex with
fuscous spots parca Ball & Beamer

17(14). Style with dorsal arm elongate; vertex without

fuscous spots rostrata Ball & Beamer

— Style without elongate dorsal arm; vertex usu-
ally with fuscous spots 18

18(17). Aedeagus with shaft shortened, widest in api-
cal 1/2, apical hook short; style with conspicu-
ous lateral flange 19

— Aedeagus with shaft longer, not widest in api-
cal 1/2, apical hook longer; style without lat-
eral flange fredonia Ball & Beamer

19(18). Aedeagus with pronounced keels on ventral
margin of shaft, shorter; hind tibial spur 1/2
1st tarsomere; from SW cursa, n. sp.

— Aedeagus without pronounced keels on shaft,
longer; hind tibial spur longer; from Califor-
nia (east face of Sierras) hemijona, n. sp.

20(6). Style with apical dorsal arm truncate; vertex
commonly with fuscous spots; plates rounded
apically incerta Ball & Beamer

— Style with apical dorsal arm rounded or acute;
vertex usually without fuscous spots; plates
variable 21

21(20). Plates rounded apically; style with ventral

apical arm thickened, well developed

kadokana KnuU

— Plates truncate apically; style with apical lat-
eral flange; ventral apical arm not as well de-
veloped; vertex occasionally with fuscous
spots salsa Ball & Beamer

22(5). Vertex usually without fuscous spots; aedea-
gus with serrated ventral margin; plates not
elongate 23

— Vertex usually with fuscous spots, or if absent,
then aedeagus without ventral serrations;
plates may be elongate 29

23(22). Style distinctly bifid apically; connective
shorter than style; aedeagus without apical
crests on shaft 25

— Style with only dorsal arm developed; con-
nective variable in length; aedeagus with
crests on shaft 24

24(23). Aedeagus with apical crests on shaft; connec-
tive equal to style in length; vertex occasion-
ally with fuscous spots

kanabana Ball & Beamer

— Aedeagus with medial crests on shaft; connec-
tive shorter than style; vertex without spots
furnaca, n. sp.

25(23). Style with apical lateral flange, apical arms
equal in length; aedeagus with shaft elongate
'. 26

— Style without flange, dorsal arm longer;
aedeagus with shaft shortened, acute apically

! 27

26(25). Style with conspicuous lateral flange; plates
short, truncate apically; vertex length less
than interocular width nmrthae, n. sp.

— Style with lateral flange smaller; plates
rounded apically, vertex length may equal
interocular width bifida Ball & Beamer

27(25). Style conspicuously bifid, dorsal arm slender,

tapered iitahna Osborn

— Style less bifid, dorsal arm robust 28

28(27). Style with dorsal arm acute apically; vertex

occasionally with spots yumana Osborn

— Style with dorsal arm bulbous apically; vertex
without spots deserta, n. sp.

29(22). Tibial spur 2/3 length of 1st tarsomere; plates
elongate, acute apically, exceeding apex of

styles; outer arm of style acute apically

inagdalena Baker

— Tibial spur variable; plates shorter; style
equal to or exceeding apex of plate, outer arm
variable 30

30(29). Style with dorsal arm acute apically 31

— Style with dorsal arm rounded or truncate . . 32

31(30). Aedeagus with shaft approximately 1 1/2X
length of dorsal apodeme, with minute serra-
tions on ventral margin . . . laeta Ball & Beamer

— Aedeagus with shaft 2X length of dorsal
apodeme, with conspicuous serrations on
ventral margin sttjlosa, n. sp.

32(30). Style with outer arm rounded; aedeagus with
shaft only slightly widened or tapered in api-
cal 1/2, ventral margin with or without serra-
tions 33

— Style with outer arm truncate; aedeagus with
shaft widest in apical 1/2, ventral margin not
serrated incongrua Baker

33(32). Aedeagus with shaft short, widest at mid-
length; style with large lateral apical flange
tenera Ball & Beamer

— Aedeagus with shaft longer, even-margined
or slightly enlarged apically; style without
conspicuous flange 34

34(33). Style deeply bifid apically, ventral arm
widened, dorsal arm even-margined; aedea-
gus with shaft slightly widened apically ....
vativala, n. sp.

— Style less bifid, with inconspicuous apical lat-
eral flange, dorsal arm tapered; aedeagus with

shaft even-margined

terebrans (Gillette & Baker)

Character.s Used in Character Code

Character]. no fuscous spots on vertex.

1 fuscous spots may be present.
Character 2. hind tibial spur 1/2 1st tarsomere or
less.
1 = spur approximate]} length of 1st tar-
somere.

1988

Blocker, Johnson: Subgenus Athysanella

character 3.

Character 4.
Characters.

Character 6.

Character 7.
Character 8.

= ocellus approximately its diameter
from eye.

1 = ocellus more remote.

--^ male plates unicolorous.

1 male plates embrowned apically.

^ male plates elongate.

1 ^ male plates rounded apically.

2 = male plates truncate apically.

= male pygofer with caudal margin
rounded.

1 = male pygofer with caudal margin ex-
tended.

^ anal tube not reaching pygofer apex.

1 = anal tube equal to or exceeding
pygofer apex.

= male pygofer not embrowned.

1 = male pygofer embrowned caudoven-

trally.
Character 9. = male plates exceeding apex of
pygofer.

1 = male plates equal to apex of pygofer.

2 = male plates not reaching apex of

pygofer.
Character 10. = styles not reaching apex of pygofer.

1 = styles equal to apex of pygofer.

2 = styles exceed apex of pygofer.
Character 11. = styles not reaching apex of plates.

1 = styles equal to apex of plates.

2 = styles exceed apex of plates.
Character 12. = styles without lateral flange.

1 = styles with lateral flange.

2 = styles with ventral arm thickened.
Character 13. = connective shorter than style.

1 = connective as long as or longer than
style.
Character 14. = aedeagus with caudal surface
smooth.
1 = aedeagus with caudal surface
troughlike.
Character 15. = shaft of aedeagus flared in caudal
aspect.
1 = shaft even-margined in caudal as-
pect.
Character 16. = aedeagus with no basal processes on
shaft.
1 = shaft with basal processes.
Character 17. = aedeagus with caudal margin of shaft
smooth.
1 = caudal margin serrate.
Character 18. = aedeagus with dorsal apodeme sim-
ple.
1 = dorsal apodeme enclosing shaft
basally.

Athysanella gardenia Osborn
Figs. 1-5

Athysanella gardenia Osborn 1930:701.
Gladionura frigida Osborn 1930:709.

Length of male 2.5 to 2.9 mm, female 3.2 to
3.4 mm; head width of male 1.0 to 1.05 mm,
female 1.05 to 1.15 mm; pronotal width of
male 0.95 to 1.05 mm, female 1.0 to 1.1 mm;
interocular width of male 0.45 to 0.55 mm.

female 0.5 to 0.55 mm; vertex length of male
0.35 to 0.4 mm, female 0.35 to 0.45 mm;
pronotal length of male 0.3 to 0.4 mm, female
0.3 to 0.35 mm. Vertex length 0.76 to 0.8
interocular width; pronotal length 0. 85 to 0. 88
vertex length.

Character code: 1-1-1-1-2-0-0-0-2-1-2-0-0-
1-0-0-0-0.

Lectotype, male (No. 43183), and lectoallo-
type, female (No. 43183), from Garden of the
Gods, Colorado, (Webster) No. 7104, in
USNM.

This species is related to pastora but can be
distinguished by the apex of the style, which is
narrowed (Fig. 2), and the posterior margin of
the plate, which is embrowned and has a me-
dian projection. The style occasionally has
macrosetae; long-winged specimens have
been observed. This species has been col-
lected in Colorado and Wyoming.

Athysanella robusta Baker

Figs. 6-9

Athysanella robusta Baker 1898:187.
Athysanella montana Osborn 1930:700.

Length of male 2.3 to 3. 1 mm, female 3.4 to
3.7 mm; head width of male 0.95 to 1. 15 mm,
female 1.05 to 1.15 mm; pronotal width of
male 0.9 to 1. 15 mm, female 1.05 to 1. 15 mm;
interocular width of male 0.4 to 0.55 mm,
female 0.5 to 0.55 mm; vertex length of male
0.35 to 0.5 mm, female 0.4 to 0.5 mm; prono-
tal length of male 0.3 to 0.4 mm, female 0.35
to 0.4 mm. Vertex length 0.8 to 0.9 interocu-
lar width; pronotal length 0.76 to 0.88 vertex
width.

Character code: 0-0-1-1-2-1-1-0-2-0-2-0-1-
0-0-1-0-0.

The type of this species has not been stud-
ied; identification is based on determined ma-
terial from various collections and the litera-
ture.

Athysanella robusta is related to pastora
but can be separated by the unique shape of
the aedeagus (Fig. 6), the pronotum nearly as
wide as the head, and the connective, which is
as long as the style. There are no fuscous spots
on the vertex of this species. It has been col-
lected in Colorado, Montana, Nebraska,
North Dakota, Wyoming, and Canada.

Athysanella pastora Blocker, n. sp.

Figs. 10-13

Length of male 2.5 to 2.7 mm, female 3.6 to

Great Basin Naturalist Memoirs

No. 12

V^^

Figs. 1-1.3. Figs. 1-5. Athysanella gardenia: 1, aedeagus, lateral view; 2, style, broad aspect; 3, pygofer, lateral view;
4, plate, ventral view; 5, female sternum VII, ventral view. Figs. 6-9. Athysanella rohusta. 6, aedeagus, lateral view;
7, style, broad aspect; 8, pygofer, lateral view; 9, female sternum VII, ventral view. Figs. 10-13. Athysanella pastora:
10, aedeagus, lateral view; 11, style, broad aspect; 12, pygofer, lateral view; 13, female sternum VII, ventral view.

3.9 mm; head width of male 1.05 to 1.15 mm,
female 1.2 to 1.25 mm; pronotal width of male
0.9 to 1.0 mm, female 1.05 to 1. 15 mm; inter-
ocular width of male 0.4 to 0.5 mm, female 0.5
to 0.6 mm; vertex length of male 0.35 to 0.45
mm, female 0. 45 to 0. 5 mm ; pronotal length of
male 0.3 to 0.4 mm, female 0.35 to 0.4 mm.
Vertex length 0.84 to 0.95 interocular width;
pronotal length 0.76 to 0.94 vertex length.

Character code: 1-1-0-1-1-1-0-1-1-2-2-0-0-
0-0-0-0-0.

Color stramineous; vertex with fuscous

spots; vertex, pronotum, and abdomen with
additional, conspicuous brown pattern; fore-
wings commonly with darkened stripes; face
with lateral fuscous marks; legs with various
amounts of fuscous coloring.

Forewings brachypterous, exposing 3 to 4
pregenital abdominal terga; ocellus approxi-
mately 1.5 its diameter from eye; hind tibial
spur 3/4 length of 1st tarsomere; female ab-
dominal sternum VII (Fig. 13) variable but
hind margin usually with conspicuous lateral
lobes, medial third truncate or rounded, vari-
ously embrowned.

1988

Blocker, Johnson: Subgenus Athysanella

Pygofer with caudal margin extended,
slightly embrowned on caudoventral border,
densely set with macrosetae (25+); anal tube
attains apex of pygofer; valve with caudal mar-
gin rounded; plates rounded and conspicu-
ously embrowned apically, slightly separated
at base; connective 2/3 or less length of style;
styles rounded apically, with conspicuous
preapical lobe, apex slightly exceeding apex of
plates; aedeagus with shaft simple, slightly
enlarged apically, elongate, with dorsal
apodeme slightly overlapping base of shaft.

Holotype, male, from Chaves Co., New
Mexico, Bitter Lake NWR, 14 Aug 1984 (R. F.
Whitcomb); 4 male and 8 female paratypes,
same data. Holotype and paratypes in USNM;
paratypes in IPL and KSU.

Athysanella pastora is related to robusta
and glohosa but can be separated from the
former by the absence of an aedeagal process.
It can be separated from glohosa by the shape
of the apex of the style, which is not widened
(Fig. 11), and by the shaft of the aedeagus
(Fig. 10), which is elongate and not serrated.
This species has been collected only on gyp
grama, Bouteloua hreviseta, in gypsum flats of
southeastern New Mexico.

Athysanella glohosa Ball & Beamer

Figs. 14-15

Athysanella glohosa Ball & Beamer 1940:18.

Length of male 2.9 mm; head width 1.2
mm; pronotal width 1.1 mm; interocular
width 0.45 mm; vertex length 0.35 mm;
pronotal length 0.4 mm. Vertex length 0.78
interocular width; pronotal length 1.07 vertex
length. Female unknown.

Character code: 1-0-1-1-2-1-1-0-1-1-1-0-0-
1-0-0-1-0.

Holotype, male, from Grand Canyon, Ari-
zona, 4 Aug 1930 (E. D. Ball) in USNM. Male
and female paratypes, same data, at KU.

Athysanella glohosa is related to gardenia
but can be separated by the shape of the shaft
of the aedeagus, which is serrate at midlength
(Fig. 14). Female paratypes examined are
doubtfully conspecific with the male. This
species is known only from its type locality in
Arizona; a specimen from Utah is assigned to
this species but may represent a new species.

Athysanella planata Ball & Beamer

Figs. 16-18

Athysanella planata Ball & Beamer 1940:24.

Length of male 2.6 to 2.7 mm, female 3.4 to
4.0 mm; head width of male 1.0 to 1.05 mm,
female 1. 1 to 1. 15 mm; pronotal width of male
0.9 to 1.0 mm, female 1.0 to 1.1 mm; interocu-
lar width of male 0.35 to 0.4 mm, female 0.4 to
0.5 mm; vertex length of male 0.35 mm to 0.4
mm, female 0.4 to 0.45 mm; pronotal length of
male 0.3 to 0.35 mm, female 0.3 to 0.35 mm.
Vertex length 0.87 to 1.0 interocular width;
pronotal length 0.8 to 0.86 vertex length.

Character code: 0-1-0-0-1-0-0-0-1-1-1-2-0-
0-0-0-0-0.

Holotype, male, and allotype, female, from
Newberry Springs, California, 30 July 1936
(D. R. Lindsay and R. H. Beamer) in KU.

Athysanella planata is related to krameri
but can be separated by the presence of pro-
cesses on the base of the shaft of the aedeagus
(Fig. 16), a more slender apical arm of the
style (Fig. 17), and the absence of fuscous
spots on the vertex. This species has been
collected only from the type locality in Cali-
fornia.

Athysanella krameri Blocker, n. sp.

Figs. 19-21

Length of male 3.0 to 3. 1 mm, female 4.4 to
4.7 mm; head width of male 1.05 to 1.15 mm,
female 1.2 to 1.25 mm; pronotal width of male
1.0 to 1.1 mm, female 1.1 to 1.2 mm; interocu-
lar width of male 0.45 to 0.5 mm, female 0.55
to 0.6 mm; vertex length of male 0.35 to 0.45
mm, female 0.45 to 0.5 mm; pronotal length of
male 0.3 to 0.35 mm, female 0.35 to 0.4 mm.
Vertex length 0.83 to 0.95 interocular width;
pronotal length 0.77 to 0.87 vertex length.

Character code: 1-1-0-0-1-1-0-0-2-1-2-0-0-
0-0-0-0-0 .

Color stramineous; vertex commonly with
fuscous spots, often with orange dashes; male
pygofer commonly with conspicuous fuscous
spot; remainder of body mostly unicolorous;
female without fuscous spots on vertex.

Forewings brachypterous, exposing 3.5 to
4.5 pregenital abdominal terga; ocellus 1.5 its
diameter (or less) from eye; hind tibial spur
approximately 3/4 length of 1st tarsomere; fe-
male abdominal sternum VII (Fig. 21) with
elongate lateral lobes and rounded medial
lobe on hind margin.

Pygofer rounded apically, with about 20
macrosetae; valve rounded on apical margin;
plates rounded apically, slightly separated or
touching at base; connective less than 3/4

Great Basin Naturalist Memoirs

No. 12

Figs. 14-27. Fig.s. 14-15. Athijsanella globosa: 14, aedeagus, lateral view; 15, style, broad aspect. Figs. 16-18.
Athysanella planata. 16, aedeagus, lateral view; 17, style, broad aspect; 18, female sternum Vll, ventral view. Figs.
19-21. Athysanella krameri: 19, aedeagus, lateral aspect; 20, style, broad aspect; 21, female sternum Vll, ventral
aspect. Figs. 22-23. Athysanella whitcombi: 22, style, broad aspect; 23, female sternum Vll, ventral view. Figs. 24-27.
Athysanella plana: 24, aedeagus, lateral view; 25, aedeagus, anterodorsal view; 26, style, broad aspect; 27, female
sternum VII, ventral view.

length of style; styles (Fig. 20) bifid apically,
dorsal arm short, robust, no preapical lobe,
exceeding apex of plates, equal to apex of
pygofer; aedeagus (Fig. 19) with shaft short,
widest basally, and tapered to acute apical
hook.

Holotype, male, from Little Lake, Inyo
Co., California, 7 June 1929 (E. P. Van
Duzee); 5 male and 6 female paratypes, same

data; 3 males from Olancha, Inyo Co., 8 June
1929; 1 male from Lone Pine, Inyo Co., 10
June 1929. Holotype and paratypes in CAS,
paratypes in KSU.

Athysanella krameri is related to planata
but can be separated by the absence of projec-
tions at the base of the aedeagus and the dorsal
arm of the style, which is shorter and more
robust. An undeveloped male specimen has

1988

Blocker, Johnson: Subgenus Athysanella

been observed. This species is known only
from the above locahties in CaHfornia.

Athysanella whitcombi Blocker, n. sp.

Figs. 22-23

Length of male 2.6 mm, female 4.0 mm;
head width of male 1.0 mm, female 1.15 to 1.2
mm; pronotal width of male 0.9 mm, female
1.0 to 1.05 mm; interocular width of male 0.4
mm, female 0.55 to 0.6 mm; vertex length of
male 0.35 to 0.4 mm, female 0.5 mm; pronotal
length of male 0.25 to 0.3 mm, female 0.45
mm. Vertex length 0.93 to 0.94 interocular
width; pronotal length 0.73 to 0.74 vertex
length.

Character code: 0-1-1-0-1-0-0-0-1-1-1-1-0-
0-0-0-0-1.

Color stramineous; vertex without fuscous
spots; body virtually unicolorous.

Forewings of male brachypterous, exposing
4.5 pregenital abdominal terga; ocellus ap-
proximately 2X its diameter from eye; hind
tibial spur about 1/2 length of 1st tarsomere;
female abdominal sternum VII with lateral
lobes, without medial lobe on caudal margin.

Pygofer rounded, with about 20 macrose-
tae; valve rounded on caudal margin; plates
rounded apically, touching at base then diver-
gent apically; connective 3/4 length of style;
styles bifid apically, dorsal arm slender, no
preapical lobe, exceeding apex of plates,
equal to apex of pygofer; aedeagus with shaft
and dorsal apodeme as plana.

Holotype, male, from Austin, Nevada, 12
Aug 1940 (D. E. Hardy) and a long- winged
female paratype, same data, in KU.

Athysanella whitcombi is related to plana
but can be separated by its smaller size, the
absence of fuscous spots on the vertex, and the
more slender dorsal apical arm of the style.
This species is named for R. F. Whitcomb,
who has made numerous contributions to this
research; it has been collected in Nevada.

Athysanella plana Ball & Beamer

Figs. 24-27

Athysanella plana Ball & Beamer 1940:13.

Length of male 3.0 to 3.3 mm, female 4.5 to
4.7 mm; head width of male 1.05 to 1.15 mm,
female 1.2 to 1.25 mm; pronotal width of male
1.0 to 1.05 mm, female 1.15 to 1.2 mm; intero-
cular width of male 0.45 to 0.5 mm, female
0.55 to 0.6 mm; vertex length of male 0.4 to
0.45 mm, female 0.45 to 0.5 mm; pronotal

length of male 0.3 to 0.4 mm, female 0.35 to
0.4 mm. Vertex length 0.89 to 0.95 interocu-
lar width; pronotal length 0.77 to 0.82 vertex
length.

Character code: 1-1-1-0-2-0-0-0-1-1-1-1-0-
0-0-0-0-0.

Holotype, male, and allotype, female, from
Califa, California, 12 June 1935 (P. W. Oman)
in USNM; paratypes in USNM.

Athysanella plana is related to aspera but
can be separated by the expanded base of the
dorsal apodeme of the aedeagus (Fig. 25) and
the longer dorsal apical arm of the style (Fig.
26). This species has been collected only from
the Central Valley (Fresno Co.) of California.

Athysanella aspera Ball & Beamer
Figs. 28-30

Athysanella aspera Ball & Beamer 1940:17.

Length of male 2.5 to 3. 1 mm, female 4.3 to
5.0 mm; head width of male 0.95 to 1. 15 mm,
female 1. 15 to 1.3 mm; pronotal width of male
0.9 to 1.1 mm, female 1.1 to 1.25 mm; interoc-
ular width of male 0.4 to 0.5 mm, female 0.45
to 0.5 mm; vertex length of male 0.35 to 0.45
mm, female 0.4 to 0.5 mm; pronotal length of
male 0.3 to 0.4 mm, female 0.35 to 0.45 mm.
Vertex length 0.8 to 0.95 interocular width;
pronotal length 0.76 to 0.93 vertex length.

Character code: 1-1-1-0-2-1-0-0-2-1-2-1-0-
0-0-0-0-0.

Holotype, male, and allotype, female, from
Mojave, California, 1 July 1931 (E. D. Ball) in
USNM.

Athysanella aspera is related to plana but
can be separated by the apex of the style,
which has a shorter dorsal arm (Fig. 29), and
the dorsal apodeme of the aedeagus, which is
not widened apically. In one specimen, the
vertex length equaled the interocular width.
The shaft of the aedeagus has an indistinct pair
of basal tubercles similar to planata. A male
specimen parasitized by Pipinculidae and
specimens with undeveloped genitalia have
been observed. This species has been col-
lected from the Mojave Desert in California.

Athysanella aphoda Blocker, n. sp.

Figs. 31-32

Length of male 3.5 mm; head width 1.2
mm; pronotal width 1.1 mm; interocular
width 0.5 mm; vertex length 0.45 mm; prono-
tal length 0.4 mm; female unknown. Vertex
0.9 interocular width; pronotal length 0.83 to

Great Basin Naturalist Memoirs

No. 12

Figs. 28-39. Figs. 28-30. Athysanella aspera: 28, aedeagus, lateral view; 29, style, broad aspect; 30, female sternum
VII, ventral view. Figs. 31-32. Athysanella aphoda: 31, aedeagus, lateral view; 32, style, broad aspect. Figs. 33-36.
Athysanella strobila: 33, aedeagus, lateral view; .34, style, broad aspect; 35, pygofer, lateral aspect; 36, female sternum
VII, ventral view. Figs. 37-.39. Athysanella supina: 37, aedeagus, lateral view; 38, style, broad aspect; 39, female
sternum VII, ventral view.

0.84 vertex length. Female unknown.

Character code: 1-0-1-0-2-0-1-0-2-1-2-2-0-
0-0-1-0-0.

Color stramineous; vertex and pronotum
with distinct, darker pattern; forewings with
dark stripes; face with distinct lateral arcs; legs
with slight fuscous coloring.

Forewings brachypterous, exposing 3.5
pregenital abdominal terga; ocellus approxi-

mately its diameter from eye; hind tibial spur
slightly shorter than 1st tarsomere.

Pygofer as strobila but larger; valve
rounded posteriorly; plates truncate apically;
styles (Fig. 32) with dorsal arm bulbous api-
cally; aedeagus (Fig. 31) with apical hook
more distinct than strobila , shaft with indis-
tinct, small process at base.

Holotype, male, from Lander Co. , Nevada,

1988

Blocker, Johnson: Subgenus Athysanella

30 mi N Austin, 24 Julv 1986 (H. D. Blocker)
in KSU.

Athysanella aphoda is closely related to and
possibly conspecific with strohila but can be
readily separated by its larger size, the bul-
bous apex of the dorsal arm of the style, and
the more robust aedeagus, which has a more
pronounced apical hook. This species is repre-
sented by the holotype from Nevada.

Athysanella strohila Blocker, n. sp.

Figs. 33-36

Length of male 2. 6 to 3. 9 mm, female 4. 2 to
5.3 mm; head width of male 1.05 to 1.3 mm,
female 1.15 to 1.35 mm; pronotal width of
male 0.95 to 1.25 mm, female 1.05 to 1.3 mm;
interocular width of male 0.4 to 0.55 mm,
female 0.45 to 0.65 mm; vertex length of male
0.4 to 0.55 mm, female 0.45 to 0.65 mm;
pronotal length of male 0.3 to 0.4 mm, female
0.3 to 0.45 mm. Vertex length 0.89 to 1.0
interocular width; pronotal length 0.7 to 0.82
vertex length.

Character code: 0-1-0-0-2-0-0-0-2-1-2-2-0-
1-0-0-0-0.

Color stramineous, occasionally patterned;
vertex without fuscous spots; abdomen occa-
sionally with some darker pattern; other body
areas occasionally with irregular, brown pat-
tern.

Forewings brachypterous, exposing 3.0 to
4.5 pregenital abdominal terga; ocellus ap-
proximately its diameter from eye; hind tibial
spur 3/4 length of 1st tarsomere; female ab-
dominal sternum VII (Fig. 36) with conspicu-
ous lateral lobes on posterior margin, with a
broadly rounded medial lobe.

Pygofer rounded on caudoventral margin,
with a distinct lobe (Fig. 35), with 15 or fewer
macrosetae; valve rounded posteriorly; plates
truncate apically, nearly touching basally,
then diverging; connective 3/4 length of style;
styles bifid (Fig. 34) apically, with ventral arm
thickened, exceeding apex of plates, equal to
apex of pygofer; aedeagus with shaft short,
widest in proximal 1/2, slightly tapered in api-
cal 1/2 to a single apical hook.

Holotype, male, from Lone Pine, Califor-
nia, 28 July 1940 (R. H. Beamer); 4 male and 3
female paratypes, same data; male and fe-
male, Fallon, Nevada, 12 Aug 1940 (L. C.
Kuitert); 2 males and 8 females. New Hum-
boldt Co., Nevada, 7 mi W Denio, 23 June
1971 (Oman); 6 males and 3 females. Eureka

Co., Nevada, W. Eureka, 25 July, 1986 (H.
D. Blocker); male and female, Lander Co.,
Nevada, 5 mi W Austin, 24 July 1986 (H. D.
Blocker); male, Owyhee Co., Idaho, 10 mi W
Bruneau, 9 July 1981 (H. D. Blocker, J. W.
Johnson); numerous specimens from Albert
Lake, Oregon, 5 mi N Valley Falls, 18 May
1969 (Oman), and 6 mi N Silver Lake, Ore-
gon, 17 May 1969 (Oman). Holotype and
paratvpes in KU; paratypes in KSU and
OrSU.

Athysanella strohila is closely related to as-
pera but can be separated by the shape of the
shaft of the aedeagus, which is more robust
and shorter, and by the dorsal apical arm of
the style, which is longer and acute apically. It
can be recognized by the unique lobe on the
caudal margin of the pygofer. An undevel-
oped female specimen has been examined.
This species has been collected in California,
Idaho, Nevada, and Oregon.

Athysanella supina Ball & Beamer

Figs. 37-39

Athysanella supina Ball & Beamer 1940:22.

Length of male 2.8 to 3.2 mm, female 4.3 to
4.6 mm; head width of male 1.0 to 1.1 mm,
female 1.15 to 1.25 mm; pronotal width of
male 1.0 to 1.05 mm, female 1.1 to 1.2 mm;
interocular width of male 0.45 to 0.5 mm,
female 0.55 to 0.6 mm; vertex length of male
0.45 to 0.55 mm, female 0.5 to 0.6 mm; prono-
tal length of male 0.35 to 0.4 mm, female 0.35
to 0.45 mm. Vertex length 1.0 to 1. 15 interoc-
ular width; pronotal length 0.68 to 0.75 vertex
length.

Character code: 1-1-1-0-1-0-0-0-2-1-2-1-0-
1-0-0-1-0.

Holotype, male, and allotype, female, from
Boca Chica, Texas, 30 May 1933 (P. W.
Oman) in USNM; paratypes in USNM and
KU.

Athysanella supina is related to fredonia
but can be separated by the shaft of the aedea-
gus, which has a shorter crest on the ventral
margin (Fig. 37), and the shape of the apex of
the style, which is more bifid (Fig. 38). This
species, described from Texas, has recently
been collected there (A. L. Hicks) from shore-
grass, Monathochloe littoralis.

Athysanella foeda Ball & Beamer

Figs. 40-43

Athysanella foeda Ball & Beamer 1940:19.

Great Basin Naturalist Memoirs

No. 12

Figs. 40-.52. Figs. 40-43. Athysanella foeda: 40, head and prothorax, dorsal \'ievv; 41, aedeagus, lateral view; 42,
style, broad aspect; 4.3, female sternum VII, ventral view. Figs. 44-46. Athysanella parca: 44, aedeagus, lateral view;
45, style, broad aspect; 46, female sternum VII, ventral view. Figs. 47-49. Athysanella rostrata: 47, aedeagus, lateral
view; 48, style, broad aspect; 49, female sternum VII, ventral aspect. Figs. 50-52. Athysanella fredonia: 50, aedeagus,
lateral view; 51, style, broad aspect; 52, female sternum VII, ventral view.

Length of male 3.4 to 3.6 mm, female 4.9 to

5.1 mm; head width of male 1.2 to 1.3 mm,
female 1.4 to 1.5 mm; pronotal width of male

1.2 to 1.3 mm, female 1.35 to 1.4 mm; interoc-
ular width of male 0.55 to 0.6 mm, female 0.65
to 0.75 mm; vertex length of male 0.5 to 0.55
mm, female 0.55 to 0.6 mm; pronotal length of

male 0.4 to 0.45 mm, female 0.4 to 0.45 mm.
Vertex length 0.83 to 0.88 interocular width;
pronotal length 0.76 to 0.85 vertex length.

Character code: 0-1-1-0-1-0-0-0-1-2-2-1-0-
1-0-0-0-1.

Holotype, male, and allotype, female, from
Laramie, Wyoming, 23 June 1935 (R. H. and

1988

Blocker, Johnson: Subgenus Athysanella

J. Beamer) in KU; paratypes in KU.

Athysanella foeda is related to parca but
can be separated by its larger size and the
presence of a conspicuous lateral lobe at the
apex of the style (Fig. 42). It can be separated
from fredonia by the shape of the apex of the
style, which is distinctly bifid. This species
has been collected in Wyoming.

Athysanella parca Ball & Beamer

Figs. 44-46

Athysanella parca Ball & Beamer 1940:15.

Length of male 2.7 to 3.4 mm, female 3.9 to
4.4 mm; head width of male 1.0 to 1.15 mm,
female 1. 15 to 1.4 mm; pronotal width of male
0.9 to 1.15 mm, female 1.05 to 1.25 mm;
interocular width of male 0.4 to 0.5 mm, fe-
male 0.5 to 0.65 mm; vertex length of male 0.4
to 0.45 mm, female 0.45 to 0.55 mm; pronotal
length of male 0.3 to 0.4 mm, female 0.4 to
0.45 mm. Vertex length 0.89 to 0.95 interocu-
lar width; pronotal length 0.82 to 0.88 vertex
length.

Character code: 1-1-0-0-1-0-0-0-2-1-2-2-0-
0-0-0-0-0.

Holotype, male, and allotype, female, from
Belen, New Mexico, 20 July 1936 (R. H.
Beamer and D. R. Lindsay) in KU; paratypes
inKU.

Athysanella parca is related to foeda but
can be separated by the absence of a lateral
flange on the style and the usual presence of
fuscous spots on the vertex. It can be sepa-
rated from incerta by the shape of the apex of
the shaft of the aedeagus, which has a single
acute apical hook (Fig. 44). A long-winged
female has been examined. This species has
been collected in Colorado and New Mexico;
in New Mexico, the host is saltgrass, Distichlis
stricta.

Athysanella rostrata Ball & Beamer

Figs. 47-49

Athysanella rostrata Ball & Beamer 1940:21.

Length of male 3.5 mm, female 5.0 to 5.1
mm; head width of male 1.15 to 1.25 mm,
female 1.3 to 1.35 mm; pronotal width of male
1.1 to 1.15 mm, female 1.2 to 1.25 mm; intero-
cular width of male 0.5 to 0.55 mm, female 0.6
to 0.65 mm; vertex length of male 0.45 to 0.5
mm, female 0.5 to 0.55 mm; pronotal length of
male 0.35 to 0.4 mm, female 0.4 to 0.45 mm.
Vertex length 0.86 to 0.91 interocular width;

pronotal length 0.75 to 0.78 vertex length.

Character code: 0-1-0-0-2-0-0-0-2-2-2-2-0-
1-0-0-0-0.

Holotype, male, and allotype, female, from
Perris, California, 5 June 1935 (P. W. Oman)
in USNM; paratypes in USNM.

Athysanella rostrata is related to foeda but
can be separated by the distinctly elongate
dorsal arm of the style (Fig. 48). A paratype
with undeveloped genitalia has been exam-
ined. This species has been collected only
from its type locality in southern California.

Athysanella fredonia Ball & Beamer

Figs. 50-52

Athtjsanella fredonia Ball and Beamer, 1940:12 (in part).

Length of male 2.6 to 2.9 mm, female 3.9 to
4.3 mm; head width of male 1.0 to 1.15 mm,
female 1. 15 to 1.4 mm; pronotal width of male
0.95 to 1.1 mm, female 1.0 to 1.25 mm; intero-
cular width of male 0.4 to 0.45 mm, female
0.45 to 0.6 mm; vertex length of male 0.35 to
0.4 mm, female 0.4 to 0.5 mm; pronotal
length of male 0.3 to 0.35 mm, female 0.35 to
0.4 mm. Vertex length 0.83 to 0.94 interocu-
lar width; pronotal length 0.75 to 0.87 vertex
length.

Character code: 1-1-0-0-1-0-0-0-1-1-1-0-0-
1-0-0-1-0.

Color stramineous; vertex with fuscous
spots anteriorly; pronotum commonly with
small brown spots; other dorsal body pattern
variable; face with lateral fuscous marks; legs
with various amounts of fuscous coloring.

Forewings brachypterous, exposing 3.5 to
4.0 pregenital abdominal terga; ocellus ap-
proximately its diameter from eye; hind tibial
spur over 3/4 length of 1st tarsomere; female
abdominal sternum VII (Fig. 52) with lateral
lobes extending posteriorly slightly more than
median lobe.

Pygofer rounded apically, densely set with
short setae; valve rounded on apical margin;
plates rounded apically, slightly separated at
base; connective shorter than style; styles
(Fig. 51) widened apically, nearly truncate,
dorsal arm acute apically, curved, usually
with conspicuous lateral flange, equal to apex
of plates and pygofer; aedeagus (Fig. 50) with
shaft not widened apically, finely serrate on
ventral margin.

Holotype, male, from Fredonia, Arizona, 6
Aug 1930 (E. D. Ball) in USNM.

Great Basin Naturalist Memoirs

No. 12

Athysanella fredonia is related to cursa but
can be separated by the longer shaft of the
aedeagus, which is not widened apically. Ex-
amination of the holotype has revealed that
the male genitalia differ from the genitalia
described by Ball and Beamer (1940). For this
reason, fredonia is redescribed and a new
species, cursa, which agrees with the pub-
lished illustration, is described. It has been
collected in Arizona, Colorado, New Mexico,
Nevada, and Utah, where it occurs on galleta,
Hilariajamesii. At the type locality, it occurs
with cursa on the same host (R. F. Whit-
comb).

Athysanella cursa Blocker, n. sp.

Figs. 53-55

Athysanella fredonia Ball & Beamer (in part).

Length of male 2.5 to 3.0 mm, female 4.0 to

4.4 mm; head width of male 1.05 to 1. 15 mm,
female 1.2 to 1.3 mm; pronotal width of male
0.95 to 1.05 mm, female 1.1 to 1.2 mm; intero-
cular width of male 0.4 to 0.45 mm, female 0.5
to 0.6 mm; vertex length of male 0.35 to 0.4
mm, female 0.45 to 0.5 mm; pronotal length of
male 0.3 to 0.35 mm, female 0.35 to 0.4 mm.
Vertex length 0.88 to 0.94 interocular width;
pronotal length 0.8 to 0.88 vertex length.

Character code: 0-1-0-0-1-0-0-0-1-1-1-1-0-
1-0-0-1-0.

Color stramineous with fuscous spots on
vertex; vertex, pronotum, and abdomen vari-
ously patterned with brown markings;
forewings commonly with brown stripes; face
with lateral fuscous marks, legs with various
amounts of fuscous coloring.

Forewings brachypterous, exposing 3.5 to

4.5 pregenital abdominal sterna; ocellus ap-
proximately 1.5X its diameter from eye; hind
tibial spur usually about 1/2 length of 1st tar-
somere; female abdominal sternum VII (Fig.
55) with lateral lobes slightly longer than me-
dian lobe.

Pygofer rounded but slightly extended api-
cally; densely set with short setae; valve with
caudal margin rounded; plates rounded api-
cally; connective shorter than styles; styles
(Fig. 54) widened apically and shallowly bifid,
dorsal arm narrowed and acute apically, usu-
ally with a conspicuous lateral flange, equal to
apex of plates and pygofer; aedeagus (Fig. 53)
with shaft short, lightly serrate on ventral
margin, with pronounced flanges on ventral
margin, slightly widest in apical 1/2.

Holotype, male, from Whites City, New
Mexico, 13 July 1936 (R. H. Beamer). One
male and two female paratypes from Flagstaff,
Arizona, 27 July 1936 (R. H. Beamer); two
males from Flagstaff, Arizona, 21 July 1936
(D. R. Lindsey); female from St. Johns, Ari-
zona, 26 July 1936 (R. H. Beamer); two males
from Estancia, New Mexico, 24 June 1940 (R.
H. Beamer); five males and one female from
Cuervo, New Mexico, 23 June 1940 (R. H.
Beamer). Holotype in KU; paratypes in KU
and KSU.

Athysanella cursa is closely related to hemi-
jona but can be separated by the shorter
aedeagus with pronounced flanges on the ven-
tral margin, the shorter tibial spurs, and the
pygofer, which is slightly extended apically.
The plates occasionally have a macroseta.
Long-winged specimens and specimens para-
sitized by Dryinidae have been observed.
This species is illustrated as fredonia in Ball
and Beamer (1940), but a study of the holo-
type of fredonia reveals that this is a different
species. It has been reported from the Desert
Plains region of Arizona, Colorado, New Mex-
ico, and Utah, where it occurs on galleta,
Hilariajamesii.

Athysanella he?nijona Blocker, n.

Figs. ,56-58

sp.

Length of male 3.2 to 3.3 mm, female 4.8 to
4.9 mm; head width of male 1.2 to 1.25 mm,
female 1.3 to 1.4 mm; pronotal width of male
1. 1 to 1. 15 mm, female 1.25 to 1.3 mm; intero-
cular width of male 0.45 to 0.55 mm, female
0.5 to 0.55 mm; vertex length of male 0.4 to
0.45 mm, female 0.5 to 0.55 mm; pronotal
length of male 0.35 to 0.4 mm, female 0.4 to
0.45 mm. Vertex length 0.85 to 0.95 interocu-
lar width; pronotal length 0.73 to 0.83 vertex
length.

Character code: 1-1-0-0-1-0-0-0-1-1-1-1-0-
1-0-0-1-1.

Color stramineous; vertex with fuscous
spots; vertex, pronotum, and abdomen with
conspicuous, dark pattern; forewings with
darkened stripes; face with lateral fuscous
marks; legs with various amounts of fuscous
coloring.

Forewings brachypterous, exposing 3.5 to
4.5 pregenital abdominal terga; ocellus ap-
proximately 1.5X its diameter from eye; hind
tibial spur over 3/4 length of 1st tarsomere;
female abdominal sternum VII (Fig. 58) with

1988

Blocker, Johnson: Subgenus Athysanella

Figs. 53-70. Figs. 53-55. Athysanella cursa: 53, aedeagus, lateral view; 54, style, broad aspect; 55, female sternum
VII, ventral view. Figs. 56-58. Athysanella hemijona. 56, aedeagus, lateral view; 57, style, broad aspect; 58, female
sternum VII, ventral view. Figs. 59-61. Athysanella incerta: 59, aedeagus, lateral view; 60, style, broad aspect; 61,
female sternum VII, ventral view. Figs. 62-65. Athysanella kadokana: 62, aedeagus, lateral view; 63, style, broad
aspect; 64, 65, female sternum VII, ventral view. Figs. 66-70. Athysanella salsa: 66, head and pronotum, dorsal view;
67, aedeagus, lateral view; 68, style, broad aspect; 69, male plate, ventral view; 70, female sternum VII, ventral view.

median lobe approximately same length as
lateral lobes.

Pygofer rounded apically, densely set with
short, stubby setae; valve with caudal margin
broadly rounded; plates broadly rounded,
slightly separated at base; connective shorter
than style; styles (Fig. 57) widened apically,
shallowly bifid, dorsal arm constricted and

curved; preapical lobe inconspicuous, with
conspicuous apical lateral lobe, extending to
apex of plates and pygofer; aedeagus (Fig. 56)
with shaft short, slightly wider apically, ser-
rated on ventral margin.

Holotype, male, from Inyo Co., California,
Westgard Pass, 14 June 1983, sweeping
grasses (R. J. Gill and D. Blocker). Two male

Great Basin Naturalist Memoirs

No. 12

and three female paratypes, same data. Holo-
type and paratype in KSU; paratypes in CDA
and KU.

Athysanella hemijona is related to ctirsa but
can be separated by the shaft of the aedeagus,
which is longer, and with the ventral margin
not as flared. This species is known only from
its type locality in California.

Athysanella mcerta Ball & Beamer

Figs. 59-61

Athysanella incerta Ball & Beamer 1940:15.

Length of male 2.7 to 3.3 mm, female 3.9 to
4.1 mm; head width of male 1.05 to 1.1 mm,
female 1. 1 to 1.25 mm; pronotal width of male
1.0 to 1. 1 mm; female 1.0 to 1. 15 mm; interoc-
ular width of male 0.45 to 0.5 mm, female 0.5
to 0.55 mm; vertex length of male 0.35 to 0.45
mm, female 0.45 to 0.5 mm; pronotal length of
male 0.3 to 0.35 mm, female 0.35 to 0.4 mm.
Vertex length 0.84 to 0.95 interocular width;
pronotal length 0.75 to 0.82 vertex length.

Character code: 1-1-0-0-1-0-0-0-2-1-2-2-0-
0-0-0-0-0.

Holotype, male, and allotype, female, from
Lamar, Colorado, 20 Aug 1936 (R. H. Beam-
er) in KU; paratypes in KU, USNM, and
CSU.

Athysanella incerta is very closely related
to kadokana but can be separated by the
shape of the apex of the style (Fig. 60), which
is not excavated as deeply, and with the dorsal
arm neither expanded apically nor with an
acute projection. Fuscous spots are some-
times present on the vertex of incerta . Long-
winged specimens have been examined. This
species has been collected in Colorado, Kan-
sas, Montana, Nebraska, New Mexico, and
Wyoming, and appears to be a specialist on
Distichlis spicata.

Athysanella kadokana Knull

Figs. 62-65

Athysanella kadokana Knull 1951:180.

Length of male 2.7 to 3.4 mm, female 4.0 to
4.4 mm; head width of male 0.95 to 1.1 mm,
female 1. 1 to 1.25 mm; pronotal width of male
0.9 to 1.05 mm, female 1.05 to 1.15 mm;
interocular width of male 0.4 to 0.5 mm, fe-
male 0.5 to 0.6 mm; vertex length of male 0.35
to 0.45 mm, female 0.45 to 0.55 mm; pronotal
length of male 0.3 to 0.35 nmi, female 0.35 to
0.4 mm. Vertex length 0.84 to 1.0 interocular

width: pronotal length 0.72 to 0.82 vertex
length.

Character code: 0-1-1-0-1-0-0-0-1-2-2-2-0-
1-0-0-0-1.

Holotvpe, male, from Kadoka, South Da-
kota, Badlands, 19 July 1950 (H. C. Severin)
atOhSU.

This species is closely related to salsa but
can be separated by the shape of the male
plate, which is rounded, and by the apex of
the style, which has a more conspicuous ven-
tral arm (Fig. 63). The shape of the posterior
margin of the female sternum VII is variable
in this species (Figs. 64, 65). Specimens para-
sitized with Pipinculidae and Dryinidae and
specimens with undeveloped genitalia have
been examined; occasionally a specimen with
faint spots on the vertex is seen. This species,
collected in Colorado, South Dakota, Mon-
tana, Nebraska, Wyoming, and Canada, ap-
pears to be a specialist on Distichlis spicata.

Athysanella salsa Ball & Beamer

Figs. 66-70

Athysanella salsa Ball & Beamer 1940:20.

Length of male 2.9 to 3.2 mm, female 3.6 to
4.4 mm; head width of male 1.0 to 1.15 mm,
female 1.0 to 1.2 mm; pronotal width of male
0.9 to 1.05 mm, female 0.95 to 1.15 mm;
interocular width of male 0.45 to 0.5 mm,
female 0.45 to 0.55 mm; vertex length of male
0.4 to 0.5 mm, female 0.45 to 0.55 mm; prono-
tal length of male 0.3 to 0.4 mm, female 0.3 to
0.4 mm. Vertex length 0.94 to 1. 1 interocular
width; pronotal length 0.72 to 0.88 vertex
length.

Character code: 0-1-1-0-2-0-0-0-2-1-2-2-0-
1-0-0-0-0.

Holotvpe, male, and allotype, female, from
St. John,' Kansas, 11 Sept 1936 (R. H. Beamer)
in KU.

Athysanella salsa is closely related to
kadokana but can be separated by the trun-
cate apices of the male plates and the less
conspicuous ventral apical arm of the style
(Fig. 68). Specimens studied from Mexico
have fuscous spots on the vertex and dark
stripes on the forewings. Long-winged fe-
males have been examined. Males and fe-
males parasitized by Dryinidae and undevel-
oped specimens have been examined. This
species has been collected in Kansas, Texas,
and Mexico, and appears to be a specialist on
Distichlis spicata.

1988

Blocker, Johnson: Subgenus Atmysanella

^-m^

\r^r^

Figs. 71-86. Figs. 71-73. Athysanella kanabana: 71, aedeagus, lateral view; 72, style, broad aspect; 73, female
sternum VII, ventral view. Figs. 74-76. Athysanella furnaca: 74, aedeagus, lateral aspect; 75, style, broad aspect; 76,
female sternum VII, ventral view. Figs. 77-79. Athysanella marthae: 11, aedeagus, lateral view; 78, style, broad
aspect; 79, female sternum VII, ventral view. Figs. 80-83. Athysanella bifida: 80, aedeagus, lateral view; 81, 82, style,
broad aspect; 83, female sternum VII, ventral view. Figs. 84-86. Athysanella utahna: 84, aedeagus, lateral view; 85,
style, broad aspect; 86, female sternum VII, ventral aspect.

Athysanella kanabana Ball & Beamer

Figs. 71-73

Athysanella kanabana Ball & Beamer 1940:20.

Length of male 3. 1 to 3.6 mm, female 4.4 to
4.7 mm; head width of male 1.1 to 1.2 mm,
female 1.25 to 1.4 mm; pronotal width of male
1.05 to 1.15 mm, female 1.2 to 1.3 mm; inter-

ocular width of male 0.5 to 0.55 mm, female
0.6 to 0.7 mm; vertex length of male 0.4 to 0.5
mm, female 0.5 to 0.55 mm; pronotal length of
male 0.35 to 0.4 mm, female 0.35 to 0.45 mm.
Vertex length 0.8 to 0.95 interocular width;
pronotal length 0.76 to 0.88 vertex length.

Character code: 0-1-1-1-1-0-0-0-2-1-2-2-0-
1-0-1-1-0.

Great Basin Naturalist Memoirs

No. 12

Holotype, male, and allotype, female, from
Kanab, Utah, 9 Aug 1936 (R. H. Beamer) in
KU; paratypes in KU and USNM.

Athysanella kanabana is related to tenera
but can be separated by the embrowned area
on the plates, the aedeagus with apical crests
on the shaft (Fig. 71), and the unique shape of
the apex of the style (Fig. 72). Occasionally,
there are fuscous spots on the vertex. Unde-
veloped specimens are represented in the
paratype series. This species has been col-
lected on Sporobolus airoides in Arizona,
Kansas, New Mexico, and Utah.

Athysanella furnaca Blocker, n. sp.

Figs. 74-76

Length of male 3.1 mm, female 4.4 to 4.7
mm; head width of male 1. 15 mm, female 1.25
to 1.3 mm; pronotal width of male 1.1 mm,
female 1.2 to 1.25 mm; interocular width of
male 0.5 mm, female 0.5 to 0.55 mm; vertex
length of male 0. 5 mm, female 0.5 to 0. 55 mm;
pronotal length of male 0.4 mm, female 0.4 to
0.45 mm. Vertex length equal to interocular
wddth; pronotal length 0.78 to 0.79 vertex
length.

Character code: 0-0-1-0-2-0-0-0-2-1-2-2-0-
1-0-0-1-1.

Color stramineous, except for dark dots on
abdomen and slight fuscous on legs.

Forewings brachypterous, exposing 3.5
pregenital abdominal terga; ocellus approxi-
mately 2X its diameter from eye; hind tibial
spur 1/2 length (or less) of 1st tarsomere; fe-
male abdominal sternum VII (Fig. 76) with
short lateral lobes and slight median projec-
tion.

Pygofer with caudal margin extended, ex-
ceeding anal tube, with approximately 25 se-
tae; valve with caudal margin rounded; plates
truncate apically, barely exceeding apex of
valve; style with ventral arm short (Fig. 75),
conspicuously flattened, dorsal arm elongate;
connective 3/4 length of pygofer; aedeagus
(Fig. 74) with conspicuous keels on shaft.

Holotype, male, from Death Valley, Cali-
fornia, 5.7 mi N Furnace Cr., 19 March 1971
(Oman); two female paratypes, same data.
Holotype and paratype in OrSU; paratype in
KSU.

Athysanella furnaca is related to kanabana
but can be distinguished by the medial crests
on the shaft of the aedeagus and the connec-
tive, which is shorter than the style. This spe-

cies is known only from the type locality in the
Mojave Desert of California.

Athysanella marthae Blocker, n. sp.

Figs. 77-79

Length of male 2.9 to 3.3 mm, female 4. 1 to
4.6 mm; head width of male 1.15 to 1.25 mm,
female 1.25 to 1.35 mm; pronotal width of
male 1.1 to 1.2 mm, female 1.1 to 1.2 mm;
interocular width of male 0.45 to 0.5 mm,
female 0.5 to 0.55 mm; vertex length of male
0.4 to 0.45 mm, female 0. 45 to 0. 5 mm; prono-
tal length of male 0.35 to 0.4 mm, female 0.4
to 0.45 mm. Vertex length 0.89 to 0.95 intero-
cular width; pronotal length 0.82 to 0.94 ver-
tex length.

Character code: 0-1-1-0-2-0-0-0-2-2-2-1-0-
1-0-0-1-1.

Color uniformly stramineous, except for ir-
regular, fuscous markings on venter and legs.

Forewings brachypterous, exposing 3 to 4.5
pregenital abdominal terga; ocellus approxi-
mately 2X its diameter from eye; hind tibial
spur 2/3 length of 1st tarsomere; female ab-
dominal sternum VII (Fig. 79) with conspicu-
ous lateral lobes on hind margin.

Pygofer broadly rounded apically, with
fewer than 25 macrosetae, exceeding apex of
plates; anal tube exceeding the apex of
pygofer; valve with caudal margin rounded;
plates short, truncate, with conspicuous mi-
crosetae; style (Fig. 78) with conspicuous lat-
eral flange and preapical lobe, deeply bifid
apically, dorsal arm truncate, ventral arm
rounded, exceeding apex of plates and
pygofer; connective 3/4 length of style; aedea-
gus (Fig. 77) with caudal margin of shaft ser-
rate, troughlike in caudal view, with incon-
spicuous process apically, dorsal apodeme
simple.

Holotype, male, from Cuatro Cienegas,
Coahuila, Mexico, 9 June 1985, (A. L. Hicks)
1266B; five male and five females paratypes,
same data. Holotype and paratypes in KSU;
paratypes in IPL.

Athysanella nuirthae is related to bifida but
can be distinguished by the conspicuous lat-
eral flange of the style and the short, truncate
plates. This species has been collected only at
the type locality in Mexico on Monanthochloe
littoralis. I name this species for my wife.

Athysanella bifida Ball & Beamer

Figs. 80-83

Athysanella bifida Ball fit Beamer 1940:23.

1988

Blocker, Johnson; Subgenus Athysanella

Length of male 2.4 to 3.0 mm, female 3. 1 to
3.7 mm; head width of male 0.9 to 1.05 mm,
female 0.95 to 1. 1 mm; pronotal width of male
0.8 to 1.0mm, female 0.9 to 1.0mm;interocu-
lar width of male 0.4 to 0.45 mm, female 0.4 to
0.5 mm; vertex length of male 0.35 to 0.45
mm, female 0.4 to 0.5 mm; pronotal length of
male 0.25 to 0.35 mm, female 0.3 to 0.35 mm.
Vertex length 0.87 to 1.0 interocular width;
pronotal length 0.7 to 0.82 vertex length.

Character code: 0-1-1-0-1-0-0-0-1-2-2-1-0-
1-0-0-1-0.

Holotype, male, and allotype, female, from
Monument, Colorado, 19 Aug 1936 (R. H.
Beamer) in KU; paratypes in KU and USNM.

Athysanella bifida is related to utahna and
ijwnana but can be separated by the elongate
shaft of the aedeagus (Fig. 80), which is not
widened at midlength. Specimens parasitized
by Dryinidae and Strepsiptera, undeveloped
specimens, and long-winged specimens have
been examined. This species has been col-
lected in Arizona, Colorado, Montana, New
Mexico, North Dakota, South Dakota, Texas,
Wyoming, and Canada. In the southern part
of its range, at least, the host is blue grama,
Bouteloua gracilis (Hicks et al. 1988).

Athysanella utahna Osborn

Figs. 84-86

Athysanella utahna Osborn 1930:705.

Length of male 2.6 to 3.3 mm, female 3.9 to
4.9 mm; head width of male 0.9 to 1.05 mm,
female 1.05 to 1.25 mm; pronotal width of
male 0.85 to 1.0 mm, female 1.0 to 1.1 mm;
interocular width of male 0.35 to 0.45 mm,
female 0.45 to 0.55 mm; vertex length of male
0.4 to 0.45 mm, female 0.45 to 0.5 mm; prono-
tal length of male 0.3 to 0.35 mm, female 0.3
to 0.4 mm. Vertex length 0.94 to 1. 13 interoc-
ular width; pronotal length 0.7 to 0.82 vertex
length.

Character code: 0-1-0-0-1-0-0-0-2-1-2-0-0-
1-0-0-1-1.

Lectotype, male, and lectoallotype, fe-
male, from Ephraim, Utah, 20 July 1914 (E.
D. Ball) in USNM.

Athysanella utahna is related to yumana
but can be separated by the more slender
apical dorsal arm and less robust ventral arm
of the style (Fig. 85). It can be separated from
bifida by the shorter shaft of the aedeagus
(Fig. 84). Rarely a specimen is seen with fus-
cous spots on the vertex; the styles occasion-

ally have a macroseta. Specimens with long
wings and undeveloped genitalia and speci-
mens parasitized by Dryinidae and Pipinculi-
dae have been examined. This species has
been collected in Arizona, California, Colo-
rado, Idaho, Nevada, Utah, Oregon, Wash-
ington, Canada, and Mexico. Reported from
saltgrass, Distichlis spicata, in Mexico and
Utah.

Athysanella yumana Osborn

Figs. 87-89

Athysanella yumana Osborn 1930:704.

Length ofmale 2.8 to 3.1 mm, female 3.8 to

4.7 mm; head width ofmale 1.05 to 1. 15 mm,
female 1. 1 to 1.25 mm; pronotal width ofmale
0.95 to 1.1 mm, female 1.0 to 1.2 mm; interoc-
ular width ofmale 0.4 to 0.45 mm, female 0.45
to 0.55 mm; vertex length ofmale 0.4 to 0.45
mm, female 0.45 to 0.50 mm; pronotal length
ofmale 0.3 to 0.35 mm, female 0.3 to 0.4 mm.
Vertex length 0.94 to 1.0 interocular width;
pronotal length 0.76 to 0.82 vertex length.

Character code: 1-1-1-0-2-0-0-0-2-1-2-2-0-
1-0-0-1-0.

Lectotype, male, and lectoallotype, fe-
male, from Yuma, Arizona (H. Osborn) in
USNM; paratypes in USNM and KU.

Athysanella yumana is related to utahna
but can be separated by the shape of the apex
of the style (Fig. 88), which has more robust
ventral and dorsal arms. Occasionally, there
are fuscous spots on the vertex. The style may
have an occasional macroseta. This species has
been reported from saltgrass, Distichlis spi-
cata; it has been collected in Arizona and Cali-
fornia.

Athysanella deserta Blocker, n. sp.

Figs. 90-92

Length ofmale 3.0 to 3.3 mm, female 4.3 to

4.8 mm; head width ofmale 1.1 to 1.2 mm,
female 1.2 to 1.3 mm; pronotal width ofmale
1.05 to 1.15 mm, female 1.15 to 1.25 mm;
interocular width of male 0.45 to 0.5 mm,
female 0.5 to 0.6 mm; head width ofmale 0.45
to 0.5 mm, female 0.5 to 0.6 mm; pronotal
length ofmale 0.35 to 0.4 mm, female 0.35 to
0.45 mm. Vertex length 0.9 to 1.0 interocular
width; pronotal length 0.75 to 0.83 vertex
length.

Character code: 0-0-1-0-2-0-1-0-2-2-2-2-0-
1-0-0-1-1.

Great Basin Naturalist Memoirs

No. 12

Figs. 87-103. Figs. 87-89. Athijsanella ijmnana: 87, aedeagiis, lateral view; 88, style, broad aspect; 89, female
sternum VII, ventral view. Figs. 90-92. Athijsanella deserta: 90, aedeagus, lateral view; 91, style, broad aspect; 92,
female sternum VII, ventral view. Figs. 93-97. Athijsanella magdalena: 93, head and pronotum, dorsal view; 94,
aedeagus, lateral view; 95, style, broad aspect; 96, male plate, ventral view; 97, female sternum VII, ventral view. Figs.
98-100. Athijsanella laeta. 98, aedeagus, lateral view; 99, style, broad aspect; 100, female sternum VII, ventral view.
Figs. 101-103. Athijsanella stijlosa: 101, aedeagus, lateral view; 102, style, broad aspect; 103, female sternum VII,
ventral view.

Color stramineous; faint orange pattern of-
ten seen on vertex; faint fuscous dots may be
present on abdominal terga; face usually with
faint lateral lines; venter with various amounts
of fuscous coloring.

Forewings brachypterous, exposing 3.5 to
4.0 pregenital abdominal terga; ocellus ap-
proximately 1.5 its diameter from eye; hind
tibial spur 1/3 to 1/2 length of 1st tarsomere;
female abdominal sternum VII (Fig. 92) with

conspicuous lateral lobes, inconspicuous me-
dial lobe rounded, with various amounts of
fuscous coloring.

Pygofer with caudal margin rounded, with
25 -h microsetae; plates separated at base,
truncate apically, not reaching apex of pygo-
fer; anal tube exceeding apex of pygofer; valve
with caudal margin rounded; connective
2/3 length of style; styles (Fig. 91) slightly
exceeding apex of pygofer with apical arm

1988

Blocker, Johnson: Subgenus Athysanella

enlarged at apex, ventral arm with small,
acute process; aedeagus (Fig. 90) with shaft
narrowed apically, finely serrate on ventral
margin, approximately 2X length of dorsal
apodeme.

Holotype, male, from Death Valley, Cali-
fornia, 36 mi S Furnace Cr., 19 March 1971
(Oman); 18 male and 15 female paratypes,
same data. Holotypes and paratypes in OrSU;
paratypes in KSU.

Athysanella deserta is related to ijiwiana
but can be separated by the shape of the apical
arms of the style with the dorsal arm enlarged
at the apex and the ventral arm with a small,
acute process. This species has been collected
only at the type locality in the Mojave Desert
of California.

Athysanella magdalena Baker

Figs. 93-97

Athysanella magdalena Baker 1898:185.

Length ofmale 2.2 to 2.7 mm, female 3.0 to
3.6 mm; head width ofmale 0.85 to 0.95 mm,
female 0.95 to 1. 1 mm; pronotal width ofmale
0.75 to 0.9 mm, female 0.9 to 1.1 mm; interoc-
ular width ofmale 0.3 to 0. 4 mm, female 0. 4 to
0.5 mm; vertex length ofmale 0.3 to 0.4 mm,
female 0.35 to 0.45 mm; pronotal length of
male 0.25 to 0.35 mm, female 0.3 to 0.35 mm.
Vertex length 0.93 to 1.1 interocular width;
pronotal length 0.8 to 0.91 vertex length.

Character code: 1-1-0-0-0-0-0-0-0-2-0-1-0-
1-0-0-1-0.

Lectotype, female, from Magdalena Mts.,
New Mexico, Aug 1894 (F. H. Snow), and
lectoallotype, male, from Forresters Ranch,
Laramie Co. , Colorado, 3 Aug 1896 (No. 2013
ofBaker)inUSNM.

Athysanella magdalena is related to tenera
but can be separated by the elongate plates
and the shape of the apex of the style (Fig. 95),
which is more clearly bifid. Plates occasionally
have one or more macrosetae, and occasion-
ally the male hind tibial spur is undeveloped.
Numerous undeveloped specimens, speci-
mens parasitized by Dryinidae and Strep-
siptera, and long-winged males and females
have been examined. This species, collected
in Arizona, California, Colorado, Montana,
New Mexico, Nevada, Oklahoma, Texas,
Utah, Wyoming, and Mexico, is a common
specialist of blue grama, Bouteloua gracilis
(Whitcomb et al. 1987, Hicks et al. 1988).

Athysanella laeta Ball & Beamer

Figs. 98-100

Athysanella laeta Ball and Beamer, 1940:16.

Length ofmale 2.7 to 2.9 mm, female 3. 8 to
4.1 mm; head width ofmale 1.0 to 1.1 mm,
female 1.1 to 1.2 mm; pronotal width ofmale
0.95 to 1.05 mm, female 1.05 to 1.15 mm;
interocular width of male 0.45 to 0.5 mm,
female 0.5 to 0.55 mm; vertex length ofmale
0.35 to 0.4 mm, female 0.4 to 0.45 mm; prono-
tal length ofmale 0.35 to 0.4 mm, female 0.35
to 0.45 mm. Vertex length 0.75 to 0.89 intero-
cular width; pronotal length 0.87 to 1.0 vertex
length.

Character code: 1-1-1-0-2-0-0-0-2-1-2-0-0-
1-0-0-1-1.

Holotype, male, from Huachuca Mts., Ari-
zona, 15 July 1934 (E. D. Ball); allotype, fe-
male, same data except 14 July, in USNM;
paratypes in USNM and KU.

Athysanella laeta is related to incongrua
but can be separated by its usually smaller size
and the shape of the shaft of the aedeagus (Fig.
98), which is shorter and not as avicephaliform
apically. Some of the male paratypes have
undeveloped genitalia. This species has been
collected in New Mexico and Arizona.

Athysanella stylosa Blocker, n. sp.

Figs. 101-103

Length ofmale 2.4 to 2.8 mm, female 3.8 to
4.1 mm; head width ofmale 0.95 to 1.1 mm,
female 1. 15 to 1.2 mm; pronotal width ofmale
0.9 to 1.0 mm, female 1.05 to 1.1 mm; interoc-
ular width ofmale 0.35 to 0.45 mm, female
0.45 to 0.5 mm; vertex length ofmale 0.35 to
0.45 mm, female 0.4 to 0.5 mm; pronotal
length ofmale 0.3 to 0.35 mm, female 0.3 to
0.4 mm. Vertex length 0.93 to 1.06 interocu-
lar width; pronotal length 0.75 to 0.82 vertex
length.

Color stramineous with brown pattern on
vertex and pronotum, vertex with fuscous
spots, wings with brown stripes, face with
brown arcs.

Forewings brachypterous, exposing 3 to 4.5
pregenital abdominal terga; ocellus approxi-
mately its diameter from eye; hind tibial spur
3/4 1st tarsomere; female abdominal sternum
VII (Fig. 103) with well-developed lateral
lobes, slightly concave medially.

Pygofer broadly rounded apically, with
fewer than 25 setae; anal tube exceeds apex of

Great Basin Naturalist Memoirs

No. 12

pygofer; valve with caudal margin rounded;
plates rounded apically; styles (Fig. 102) bifid,
dorsal arm acute, ventral arm truncate, with-
out lateral flange or preapical lobe, exceeding
apex of plates and pygofer; connective 3/4
length of style; aedeagus (Fig. 101) with shaft
serrate on caudal margin, troughlike in caudal
view, with inconspicuous process apically.

Holotype, male, from Roswell, Chaves
Co., New Mexico, Bitter Lake Natl. WLR, 14
Aug 1984 (R. F. Whitcomb) 0868A; 17 male
and 7 female paratypes, same data; 1 male
paratype, same data except 21 Aug 1985,
01931. Holotype and paratypes in KSU;
paratypes in USNM and IPL.

Athysanella stylosa is related to supina but
can be distinguished by the fuscous spots on
the vertex and by the shaft of the aedeagus,
which is not as widened medially. This species
is different in that the vertex length is equal to
the interocular width and fuscous spots are
present. It has been collected on gyp drop-
seed, Sporobolus nealleiji, in the gypsum flats
of southeastern New Mexico.

Athysanella incongrua Baker

Figs. 104-106

Athysanella incongrua Baker 1898:188.

Length of male 3.0 to 3.4 mm, female 4.2 to
4.8 mm; head width of male 1.1 to 1.2 mm,
female 1.15 to 1.35 mm; pronotal width of
male 1.05 to 1. 15 mm, female 1. 1 to 1.25 mm,
interocular width of male 0.45 to 0.5 mm,
female 0.55 to 0.65 mm; vertex length of male
0.35 to 0.45 mm, female 0.45 to 0.5 mm;
pronotal length of male 0.35 to 0.4 mm, fe-
male 0.35 to 0.45 mm. Vertex length 0.76 to
0.85 interocular width; pronotal length 0.88 to
1.0 vertex length.

Character code: 1-1-0-0-2-1-0-0-2-1-2-2-0-
1-0-0-0-0.

Lectotype, male, from Fort Collins, Colo-
rado, 21 July (C. F. Baker) in USNM.

Athysanella incongrua is related to laeta
and terebrans but can be separated by the
length and shape of the apex of the shaft of the
aedeagus (Fig. 104). The fuscous spots on the
vertex are commonly missing. Long-winged
males and females have been examined. Spec-
imens, collected in Colorado, Iowa, Kansas,
Nebraska, New Hampshire, New Mexico,
Oklahoma, South Dakota, Texas, and Wyo-
ming, appear to be associated with little

bluestem, Schizachyrium scroparium. The
specimen from New Hampshire was collected
by P. W. Oman at Wiley House, 20 Aug 1934.

Athysanella tenera Ball & Beamer

Figs. 107-109

Athysanella tenera Ball & Beamer 1940:17.

Length of male 2.5 to 3.0 mm, female 3.9 to
4.0 mm; head width of male 1.0 to 1.1 mm,
female 1.1 to 1.2 mm; pronotal width of male
0.9 to 1.0 mm, female 1.0 to 1.15 mm; interoc-
ular width of male 0.4 to 0.45 mm, female 0.45
to 0.5 mm; vertex length of male 0.35 to 0.45
mm, female 0.4 to 0.45 mm; pronotal length of
male 0.3 to 0.35 mm, female 0.35 to 0.4 mm.
Vertex length 0.94 to 1.0 interocular width;
pronotal length 0.76 to 0.8 vertex length.

Character code: 1-1-0-0-1-0-0-0-1-0-2-2-0-
1-0-0-1-0.

Holotype, male, and allotype, female, from
Las Vegas, Nevada, 8 Aug 1936 (R. H.
Beamer) in KU; paratypes in KU.

Athysanella tenera is related to magdalena
but can be distinguished by the shorter plates
and the irregular shape of the outer apical arm
of the style (Fig. 108), which is widened and
appears to be trifid. This species is known only
from the type locality in Nevada.

Athysanella vativala Blocker, n. sp.

Figs. 110-112

Length of male 2.9 to 3.2 mm, female 4.2 to
4.6 mm; head width of male 1.05 to 1.15 mm,
female 1.25 to 1.35 mm; pronotal width of
male 1.0 to 1.15 mm, female 1.15 to 1.3 mm;
interocular width of male 0.4 to 0.55 mm,
female 0.55 to 0.65 mm; vertex length of male
0.35 to 0.5 mm, female 0.45 to 0.55 mm;
pronotal length of male 0.35 to 0.4 mm, fe-
male 0.35 to 0.45 mm. Vertex length 0.78 to
0.9 interocular width; pronotal length 0.74 to
0.94 vertex length.

Character code: 1-0-0-0-1-0-0-0-1-1-1-0-0-
1-0-0-1-0.

Color stramineous; vertex with fuscous
spots; vertex, pronotum, and abdomen with
additional brown pattern; fore wings with
darkened stripes; face with lateral fuscous
marks; legs with fuscous areas.

Forewingsbrachypterous, exposing 4 to 4. 5
pregenital abdominal terga; ocellus 2X its di-
ameter from eye or less; hind tibial spur 1/2
length of 1st tarsomere; female abdominal

1988

Blocker, Johnson: Subgenus Athysanella

113

114

Figs. 104-117. Figs. 104-106. Athysanella incongrua: 104, aedeagus, lateral view; 105, style, broad aspect; 106,
female sternum VII, ventral view. Figs. 107-109. Athysanella tenera: 107, aedeagus, lateral view; 108, style, broad
aspect; 109, female sternum VII, ventral view. Figs. 110-112. Athysanella vativala: 110, aedeagus, lateral view; 111,
style, broad aspect; 112, female sternum VII, ventral view. Figs. 113-117. Athysanella terebrans: 113, head and
pronotum, dorsal view; 114, aedeagus, lateral view; 115, 116, style, broad aspect; 117, female sternum VII, ventral

sternum VII (Fig. 1 12) with lateral lobes much
longer than medial lobe.

Pygofer rounded apically, fewer than 25
macrosetae; valve with caudal margin angu-
late; plates separated at base, rounded api-
cally; connective 3/4 length of style; styles
(Fig. Ill) deeply bifid, dorsal arm elongate
and slender, ventral arm thickened, no
preapical lobe present; aedeagus, in lateral
view, with shaft slightly widened in apical half
(Fig. 110), troughlike on ventral margin, and
slightly serrate.

Holotype, male, from Sheridan Co., Ne-
braska, 7 mi N Rushville, Hwy 87, 9 Aug 1979
(H. D. Blocker and R. A. Sweet); female
paratype, same data; 4 male and 7 female
paratypes, Cherry Co., Nebraska, S Valen-
tine, Hwy 83, mi 202, 10 Aug 1979 (H. D.
Blocker and R. A. Sweet); 2 males. Cherry
Co., Nebraska, near Manker, Hwy 20, 9 Aug
1979 (H. D. Blocker and R. A. Sweet); 1 male.
Bowman, North Dakota, 5 July 1968, GL 819
(Harris and Cooley); 2 males and 10 females,
Tryon, McPherson Co., Nebraska, 7 Aug

Great Basin Naturalist Memoirs

No. 12

1977, (R. F. Whitcomb) 0480. Holotype and
paratypes in KSU; paratypes in CNC, IPL,
USNM.

Athysanella vativala is very closely related
and possibly conspecific with terebrans but
can be separated by the shape of the apex of
the style, which has a more slender dorsal
arm, a more patterned and longer vertex, and
the ocelli usually more distant from the eye.
Undeveloped specimens and specimens para-
sitized with Strepsiptera have been exam-
ined. This species has been collected in Ne-
braska and North Dakota on Calanovilfa
longifolia.

Athysanella terebrans (Gillette & Baker)

Figs. 113-117

Euttetix terebrans Gillette & Baker 1895:102.

Length of male 2.7 to 3. 1 mm, female 4.2 to
4.6 mm; head width of male 1.0 to 1.15 mm,
female 1.2 to 1.35 mm; pronotal width of male
0.95 to 1.05 mm, female 1.15 to 1.3 mm;
interocular width of male 0.45 to 0.5 mm,
female 0.55 to 0.6 mm; vertex length of male
0.35 to 0.45 mm, female 0.45 to 0.5 mm;
pronotal length of male 0.3 to 0.4 mm, female
0.4 to 0.45 mm. Vertex length 0.75 to 0.89
interocular width; pronotal length 0.82 to 1.0
vertex length.

Character code: 1-0-0-0-1-0-0-0-1-1-1-0-0-
1-0-0-1-0.

Holotype, female, from North Park, Colo-
rado, 30 July (Gillette) in USNM; allotype,
male, from Wray, Colorado, 13 July 1899, at
Colorado State University.

Athysanella terebrans is related to incon-
grua but can be distinguished by the dorsal
apical arm of the style (Figs. 115, 116), which
is rounded, and by the shaft of the aedeagus
(Fig. 114), which is not conspicuously
widened apically; the female abdominal ster-
num VII (Fig. 117) may be variable in shape.
Long-winged and undeveloped specimens
were commonly examined. This species has
been collected in Colorado, Montana, Ne-
braska, North Dakota, South Dakota, Utah,
Wyoming, and Canada. It is reported from
Dystichlis striata in Manitoba.

Phylogeny

Phylogeny of the subgenera oi Athysanella
is discussed in Blocker and Johnson (1988).
The subgenus Athysanella is designated a sis-

ter o{ Gladionura and can be separated by the
absence of a pygofer process in the former.
The presence of a pygofer process is a conver-
gent character that occurs in two other less
closely related subgenera. We selected 40
characters that show differences within the
subgenus for analysis. An intuitive phylogeny
based on the proposed sequence of occur-
rence of major derived characters was gener-
ated, as well as an analysis by PAUP (Swofford
1986). Results of these analyses are shown in
Figures 118 and 119. PAUP generated five
equally most parsimonious trees; the tree
with no unresolved trifurcations is presented
here. A hypothetical ancestor consisting of a
uniform plesiomorphic character set was uti-
lized. Character sequences are contained in
the preceding descriptions; the entire PAUP
data set is available on request. Overall, the
PAUP cladogram and the intuitive phylogeny
are in good agreement. Species groups 1, 3, 4,
and 7 are consistently grouped together (Fig.
119), whereas the rostrata and incongrua-
planata groups are widely fragmented. It is
possible that this fragmentation of the groups
is indicative of the high degree of homoplasy
in the genus.

The robusta group is characterized by re-
tention of the rounded shape of the style apex;
the male plates are embrowned in all species
(2'); the styles are widened and often bifid
apically in all other species (1'). This group is
recognized in the PAUP output.

The rostrata group has an extended apex of
the male style (1"); PAUP has the species in
this group occurring close together but none
as sisters.

The incerta group is characterized by the
presence of a semicircular apex of the shaft of
the aedeagus (3'). There is fairly good congru-
ence in the cladogram, which includes
planata as a sister o( incerta .

The terebrans-bifida group is characterized
by an aedeagal shaft that is serrate on the
caudal margin (4'). The terebrans group has a
shortened vertex (5'); five of the seven species
are recognized by PAUP; the other two (laeta
and marthae) split out sooner but are close.
The bifida group retains an elongate vertex;
they are represented in the last half of the
cladogram, close together, but only bifida and
magdalena arc sisters.

The incongrua-planata group retains an
aedeagus with a simple shaft. PAUP separates

1988

Blocker. Johnson: Subgenus Athysanella

pastora

robust a

glob OS a

gardenia

kadokana

salsa

incerta

vativala

terebrans

lacta

marthae

fredonla

cursa

hemijona

bifida

utahna

■ yumana

■ deserta
•stylos a

■ tenera

■ magdalena
• supina

foeda

parca

incongrua

aphoda

strobila

aspera

plana

whitcombi

krameri

planata

rostrata

kanabana

furnaca

o~- en 4^ OJ t^

CQ CQ OQ OQ CO Ua

-! '-J "1 "1 ^ ■-!

o o o o o o

c c; c c c c

"X3 "1:3 13 'T3 "O "O

•^ -

■^ -

^ ■
CO -
^ -
CO
CO

I to —

ro
en
c;i

O) -

Lcs:

^ -

Oi
4^-

Oulgroup
robusla
pastora
globosa
gardenia
krameri
aspera
wbitcoinbi
plana
parca
aphoda
incerta
planata
salsa
kadokana
strobila
rostrata
incongrua
kanabana
deserta
furnaca
yumana
tenera
supina
foeda
marthae
bifida
magdalena
laeta
utahna
•stylata
• vativala
terebrans
fredonia
cursa
hemijona

Fig. 118. Intuitive cladogram.

the incongrua group to a greater degree than
any other group in the intuitive cladogram.
The planata group species are characterized
by an ocellus that is more remote from the eye
than other groups (6') and all split out in the
first half of the PAUP cladogram. Four species
{krameri, aspera, whitcombi, and p/ona) arise
from the cladogram in sequence.

Acknowledgments

Paul Oman has always been gracious and
very helpful in his support of leafhopper sys-
tematics; it is a high honor to contribute to a

Fig. 119. PAUP cladogram.

volume dedicated to his many contributions
to our science. Merv Nielson collected with
the senior author in Mexico and elsewhere
and has offered assistance throughout this
study. Likewise, the assistance of Bob Whit-
comb and Andy Hicks has added significantly,
particularly with host information and species
distribution. Bob Brooks frequently furnished
work space in the Snow Entomological Mu-
seum. Paul Cwikla has read drafts of this and
related manuscripts, and JefiFry Ediger, a stu-
dent at KSU, inked the illustrations of the
genitalia.

Great Basin Naturalist Memoirs

No. 12

Literature Cited

Ball, E. D , and R H Beamer 1940. A revision of the
genus Athysanella and some related genera (Ho-
moptera: Cicadellidae). Univ. Kansas Sci. Bull.
26: 5-82.

Blocker, H. D. 1984. Morphological irregularities in the
external genital structures of Athysanella (Ho-
moptera: Cicadellidae: Deltocephalinae). Bull.
Soc. Entomol. Suisse 57: 412.

Blocker, H. D., and J. W Johnson. 1988. Subgenera of
the genus Athysanella Baker (Homoptera: Ci-
cadellidae: Deltocephalinae) and a proposed phy-
logeny. Proc. VI Auchenorrhyncha Mtg., Torino,
Italy, September 1987.

Blocker, H. D., andC S. Wesley 1985. Distribution of
Athysanella (Homoptera: Cicadellidae: Delto-
cephalinae) in Canada and Alaska with descrip-
tions of three new species. J. Kansas Entomol.

Soc. 52: 377-386.

Hicks, A L , R F Whitcomb, H D Blocker, and K. A
Allred 1988. Why are there so many species of
Athysanella? Proc. VI Auchenorrhyncha Mtg.,
Torino, Italy, September 1987.

Swofford, D L. 1986. PAUP, phylogenetic analysis us-
ing parsimony. Private publication.

Wesley, C S , and H D Blocker 1985. Athysanella of
Mexico including descriptions of a new subgenus
and six new species (Homoptera: Cicadellidae).
Entomography 3: 163-180.

Whitcomb, R F , A L Hicks, D E Lynn, K A Allred,
andH D Blocker. 1987. Geographic variation in
host relationships of leafhoppers (Homoptera: Ci-
cadellidae) in North American Grasslands. In: M.
R. Wilson and L. R, Nault, eds., Proc. 2nd Int.
Workshop on Leafhoppers and Planthoppers of
Economic Importance. Commonwealth Inst, of
Entomol., London.

A NEW GENUS, ILEOPELTUS, RELATED TO CHLOROTETTIX
(HOMOPTERA: CICADELLIDAE)

Paul S. Cwikla'

Abstract — On the basis of their unique genitahc characters, the Neotropical leafhopper species previously treated
in the genus Doleranus Ball are placed in a new genus, Ileopeltus. Twelve species are treated, including six described
as new. A key to the species and a cladistic hypothesis using Chlorotettix Van Duzee as the outgroup are presented. The
new species of Ileopeltus include: nanocanthus (Panama), dorsalus and clavatus (Brazil and Venezuela), ventriculus
and haplus (Brazil), and blockeri (Venezuela).

This paper is the first in a series of revisions
deahng with the New World deltocephahne
genera, which have their crown completely
microsculptured. These leaflioppers, com-
monly called the broad-headed leaflioppers,
are best known from the genus Chlorotettix
Van Duzee. Species of this genus are com-
monly found in low-lying, grassy habitats of
North and South America.

Traditionally, those leaflioppers with a
completely microsculptured crown have been
restricted to two genera, Chlorotettix Van
Duzee and Doleranus Ball (Oman 1949). Lin-
navuori (1959) mentioned that some species of
Paratanus Young and Stirellus Osborn & Ball
also have their crown microsculptured, al-
though no degree of phylogenetic relationship
was implied.

More recently, I (Cwikla 1988) examined
the North American species o{ Doleranus and
found that they fell within an acceptable range
of variation for Chlorotettix. I did not, how-
ever, consider the Neotropical species of Dol-
eranus congeneric with Chlorotettix because
of the following synapomorphies: the asym-
metrical aedeagus, the aedeagus without pro-
cesses, and, with the exception of only two
species, the male plates fused with the valve.
Because of these unique attributes, the
Neotropical species of Doleranus treated by
Linnavuori (1959) and six species described as
new are designated a new genus, Ileopeltus.

Virtually nothing is known about the biol-
ogy of Ileopeltus species. Ileopeltus tethys has
been collected from sugar cane, weeds, sweet
potato (Wolcott 1923), and grassy pastures
(Caldwell and Martorell 1950) in Puerto Rico.

Hosts for the other species of Ileopeltus are
probably grasses.

With the exception of/, tethys, specimens
of Ileopeltus species are rare in collections.
For the most part they appear restricted be-
tween the Tropic of Cancer and the Tropic of
Capricorn. Specimens have been collected in
low- to mid-altitude areas.

Twelve species are included in this new
genus, of which six are described as new. In
addition, this paper provides a key to the spe-
cies and a discussion of the phylogenetic rela-
tionships among the species. Specimens were
prepared for SEM study as described in
Cwikla and Freytag (1983). Names of institu-
tions associated with abbreviations used in the
text are in the acknowledgments section.

Ileopeltus, n. gen.

Diagnosis. — Ileopeltus species can be sep-
arated from other deltocephaline genera by
the asymmetrical aedeagus without processes
and the crown completely microsculptured.

Color. — Yellowish green or ochraceous.
Markings occasionally present on crown and
forewing.

Structural features. — Small, delto-
cephaline leaflioppers. Crown roundly pro-
duced, median length slightly longer than
length next to eye. Clypellus not constructed
proximally. Forewing with cross-vein in claval
area.

Male genitalia. — Pygofer roundly pro-
duced or truncated, process present or ab-
sent. Anal tube weakly sclerotized dorsally.
Valve acutely triangular, usually fused to plate
posteriorly. Plate short or slightly longer than

'Department of Entomology, University of Missouri, Columbia. Missouri 65211, Present address: 14560 Josephine Street, Irwin, Pennsylvania 15642.

Great Basin Naturalist Memoirs
I

No. 12

Figs. 1-4. Ileopeltus tethijs Van Duzee; 1, pygofer, left lateral aspect; 2, right style, dorsal aspect; 3, aedeagus,
ventral aspect; 4, aedeagus and apex of connective, left lateral aspect.

pygofer, rounded apically, macrosetae unise-
riate, second short row of microsetaelike
structures occasionally present, lateral mar-
gin sinuate, straight, or slightly convex. Stylar
apex linear, preapical angle usually absent.
Connective as long as or shorter than aedea-
gus. Aedeagus without processes, asymmetri-
cal, apex acute in posterior aspect. Gonopore
on right side at base of shaft.

Type species. — Chlorotettix tethijs Van
Duzee 1907:71.

Distribution. — Neotropical, usually re-
stricted between the Tropic of Cancer and the
Tropic of Capricorn.

Notes. — This genus has been previously
described as Doleranus by Linnavuori (1959).
The Neotropical species are not considered
congeneric with the Nearctic forms and have
been treated as synonyms oi Chlorotettix (in-
cluding the type species of Doleranus, Tham-
notettix longulus Gillette & Baker) in a disser-
tation by Cwikla (1988).

Key to the Male Ileopeltus Species

1. Pygofer without a process (Fig. 1), or if

present then e.xtremely small (Fig. 7) 2

— Pygofer with long, distinct processes (Fig. 43) 3

2(1). Pygofer without a process (Fig. 1)

tethys (Van Duzee)

— Pygofer with short process on posterodorsal mar-
gin in lateral aspect nanocanthus , n. sp.

.3(1). Stylar apex curved laterally (Fig. 52)

Cyclops (Linnavuori)

— Stylar apex linear (Figs. 12, 30) 4

4(3). Pygofer with process directed ventrad in lat-
eral aspect ventriculus. n. sp.

— Pygofer with process directed mesad or dor-
sad in lateral aspect 5

5(4). Pygofer with process heavily sclerotized, in-
serted on medial side in lateral aspect (Fig.
27) 8

— Pygofer with process not heavily sclerotized,
not inserted on medial side in lateral aspect

(Fig. 11) 6

6(5). Aedeagal shaft wide in ventral aspect (Fig. 25)

hapliis, n. sp.

— Aedeagal shaft narrow in ventral aspect (Fig.
22) . . .' 7

7(6). Stylar apex truncate (Fig. 12) . . dorsalus, n. sp.

— Stylar apex narrow (Fig. 20) . spinosus (DeLong)

8(5). Stylar apex truncate and constricted subapi-

cally (Fig. 40) clavaUis, n. sp.

— Stylar apex rounded or if truncate, then not
constricted subapically 9

9(8). Pygofer with process inserted on middle of

ventral margin (Fig. 33) blockeri, n. sp.

— Pygofer with process inserted on posteroven-
tral margin (Figs. 27, 49) 10

10(9). Aedeagus wide and highly asymmetrical in

1988

CwiKLA: The Genus Ileopeltus

Fig. 5. Ileopeltus tethys Van Duzee, dorsal habitus.

ventral aspect (Fig. 49)

cuneus (DeLong & Martinson)

— Aedeagus narrow in ventral aspect 11

11(10). Aedeagus with large gonopore in lateral as-
pect (Fig. 32); plate fused with valve

hasttilus (DeLong & Linnavuori)

— Aedeagus with small gonopore in lateral as-
pect (Fig. 46); plate not appearing fused with
valve aberrans (Osborn)

Ileopeltus tethys (Van Duzee), n. comb.
Figs. 1-5, 57, 59; Map 1

Chlorotettix tethys Van Duzee 1907:71.

Chlorotettix hidentatus DeLong 1923:264, Wolcott 1936:

86.
Chlorotettix dilutus Oshorn 1923:73, Osborn 1935:118.
Doleranus kinonanus Ball 1936:432, Linnavuori 1959:

274-275.

Diagnosis. — Ileopeltus tethys is near /.
nanocantus and can be separated from it and
other Ileopeltus species by the lack of a
pygoferal spine.

Length. — Male 3.8-4.8 mm, female 4.1-
5.4 mm.

Color, — Yellowish green. Eye reddish
brown. Forewing yellowish subhyaline, small
brown patch occasionally present on middle of
wing and/or on claval area.

Male genitalia. — Pygofer with posterior
margin roundly produced, processes absent.
Plate short, approximately half the length of
pygofer, lateral margin straight or slightly
concave, not fused to valve. Stylar apex stout,
linear, preapical angle absent. Aedeagus
asymmetrical in posterior aspect, left subapi-
cal margin sinuate in posterior aspect, right
subapical margin straight or somewhat convex
in posterior aspect. Gonopore at base on right
side.

Female seventh sternum. — Posterior
margin shallowly excavated, median of exca-
vation with short, produced projection bear-
ing two short teeth (see Linnavuori 1959: Fig.
113g). Depth of excavation varies from only
slightly to one-third width of segment.

Type. — A single female cotype bearing the
labels "Martinique W.I., VII-26" and "Aug.
Busck collector" was examined from the
USNM.

Distribution. — ^Many specimens from the
following localities were examined: Central
America: Honduras, Mexico (Chiapas, Guer-
rero, Michoacan, Oaxaca, Tamaulipas, Vera-
cruz), Nicaragua, Panama. West Indies: An-
tigua, Cayman Islands, Dominican Republic,
Guadeloupe, Haiti, Jamaica, Montserrat,
Nevis, Puerto Rico, Trinidad, and Tobago.
South America: Venezuela (Guarico).

Specimens were collected from May
through December and are deposited in the
UPB, UCV, UKC, USNM, OSUC, and
BMNH collections.

Note. — The types of hidentatus, dilutus,
and kinonanus were not available for study;
consequently, previous synonymies of these
names under /. tethys were not verified. Be-
cause the types are females, it is doubtful that
they can be correctly associated with males at
this time.

Ileopeltus nanocanthus, n. sp.

Figs. 6-10, 58, 60; Map 2

Diagnosis. — Ileopeltus nanocanthus is
near /. tethys and can be separated from it and
other Ileopeltus species by the short pygoferal
spine.

Length. — Male 4.6-5.4 mm, female 4.8-
5.3 mm.

Great Basin Naturalist Memoirs

No. 12

Map 1. Distribution of Ileopeltiis tethijs.

Color. — Ochraceous or greenish yellow.
Eye reddish brown. Crown occasionally with
faint orange, median band. Pronotum with
four longitudinal, faint orange bands. Fore-
wing yellowish subhyaline, brown spots
present at bases of anteapical cells and on
claval area.

Male genitalia. — Pygofer with ventral
margin roundly produced into short spine on
dorsoposterior margin, spine directed medi-
ally in ventral aspect. Plate short, anterior
margin convex. Stylar apex narrowed, di-
rected posteriorly, preapical angle absent.
Aedeagus linear, asymmetrical in posterior
aspect, apex with opposite curve compared to
/. tethys. Gonopore at base on right side.

Female seventh sternum. — Posterior
margin with U-shaped excavation extending
half length of segment, base of excavation with
small V-shaped notch, notch surrounded by
light brown coloration, lateral angle rounded.

Type. — Male holotype, Panama, Canal
Zone, Curundu, 26-3'()-XI-1986, B. Sieber-
glied collector. Four female paratypes, same

data as holotype; one male paratype, Panama,
Chiriqui, Rio Colorado, 1,200 m, 8°5'N,
82°43'W, 26-XII-1974, H. Wolda collector;
male paratvpe, Panama, Panama, Las Cum-
bres, 17-23-11-1982, H. Wolda collector.
Holotype, two paratypes with same data as
holotype, and one paratype from Las Cum-
bres deposited in OSUC, remainder in HWC.
Etymology. — Greek, nano (dwarf) and
acantha (spine) refer to the small pygofer pro-

Ileopeltiis dorsalus, n. sp.

Fig.s. 11-14; Map 2

Diagnosis. — Ileopeltus dorsalus is near 7.
ventriculus and can be separated from it by
the acute, dorsally directed pygoferal spine.

LENCiTH. — Male 4.4 mm, female unknown.

Color. — Ochraceous, either without dark
markings or crown with light brown surround-
ing coronal sulcus. Eye red. Pronotum with
six light brown, longitudinal bands. Scutel-
lum with brown lateral angles. Forewing sub-
hyaline with brown patches at base of anal

1988

CwiKLA: The Genus Ileopeltus

Figs. 6-10. Ileopeltus nanocanthus, n. sp.: 6, pygofer, left lateral aspect; 7, pygofer, ventral aspect; 8, right style,
dorsal aspect; 9, aedeagus, ventral aspect; 10, aedeagus and apex of connective, left lateral aspect.

Map 2. Distribution of Ileopeltus nanocanthus (squares), /. dorsalus (circles), and I. ventriculus (triangles)

Great Basin Naturalist Memoirs

No. 12

Figs. 11-14. Ileopeltus dorsalus, n. sp.: 11, pygofer, left lateral aspect; 12, right style, dorsal aspect; 1.3, aedeagus,
ventral aspect; 14, aedeagus and apex of connective, left lateral aspect.

veins, proximal and distal ends of inner ante-
apical cell, and proximal end of central ante-
apical cell.

Male genitalia. — Pygofer with posterior
margin produced into dorsally directed spine,
acute apically, not heavily sclerotized, small
group of microsetaelike structures near mid-
dle of posterior margin. Plate triangular, lat-
eral margin straight. Stylar apex truncate, di-
rected posteriorly, preapical angle small.
Aedeagus only slightly asymmetrical.

Type. — Male holotype, Venezuela, Aragua,
El Limon, 22-11-1973, Malaise trap, C. J.
Resales collector. Male paratype, Brazil,
Ceara, Barbalha, V-1969, M. Alvarenga, B.M.
1971-165. Holotype deposited in OSUC,
paratype in BMNH.

Etymology, — Latin, dorso (back) refers to
the dorsally directed pygoferal spine.

Ileopeltus ventriciilus , n. sp.

Figs. 1.5-18; Map 2

Diagnosis. — Ileopeltus ventriculus is near
spinosus and can be separated from it and
other Ileopeltus species by the pygoferal pro-
cess directed ventrally.

Length. — Male 4.4 mm, female unknown.

Color.— Ochraceous, without distinct dark

markings. Eye reddish brown.

Male genitalia. — Pygofer with short, ven-
trally directed process inserted on posterior
margin. Plate elongate, lateral margin sinu-
ate, apex curved dorsally, bluntly rounded.
Style elongate, apex linear, slightly curved
laterally. Aedeagus elongate, broadly curved
dorsally in lateral aspect, apex acute in poste-
rior aspect.

Type. — Male holotype, Brazil, Ceara,
Crato, May 1969, M. Alvarenga collector.
Type deposited in BMNH.

Etymology. — Latin, ventricul (belly)
refers to the ventrally directed pygofer pro-
cess.

Ileopeltus spinosus (Dehong), n. comb.

Figs. 19-22; Map 3

Chlorotettix spinosus DeLong 1945:10-11.

Diagnosis. — Ileopeltus spiiiosus is near /.
dorsalus and can be separated from it and
other Ileopeltus species by the presence of a
preapical angle on the style and the unique
pygoferal process.

Length. — Male 5.2-5.5 mm, female un-
known.

Color. — Greenish yellow, without dark-
markings. Eye gray or greenish yellow.

1988

CwiKLA: The Genus Ileopeltus

Figs. 15-18. Ileopeltus ventricuhis , n. sp.: 15, pygofer, left lateral aspect; 16, right style, dorsal aspect; 17, aedeagus,
ventral aspect; 18, aedeagus and apex of connective, left lateral aspect.

Male genitalia. — Pygofer with posterior
margin broadly rounded apically, heavily
sclerotized process inserted on posteroventral
margin, directed posteriorly then curved me-
dially. Plate with lateral margin slightly sinu-
ate, apex elongate, pointed. Stylar apex nar-
rowly elongate, directed slightly laterally,
preapical angle roundly triangular. Aedeagus
elongate, not strongly asymmetrical, narrow
in ventral aspect. Gonopore on right side near
base.

Type. — Male holotype bearing the labels
"Buena Vista, Gro. [Guerrero, Mexico], 23-X-
1941, 3,400 ft." and "DeLong, Good, Cald-
well and Plummer" was examined from
OSUC.

Distribution. — Besides the type, three
paratypes from Mazaclan, Guerrero, Mexico,
3-X-1945, were examined from OSUC.

Note. — DeLong (1945) reported this spe-
cies from meadow grasses.

Ileopeltus haplus, n. sp.

Figs 23-26; Map 3

Diagnosis. — Ileopeltus haplus is near /.
spinosus and can be separated from it and
other Ileopeltus species by the unique pygofer
process and the wide aedeagus in ventral as-
pect.

Length. — Male 5.3 mm, female unknown.

Color. — Yellowish green, without dark
markings. Eye yellowish green.

Male genitalia. — Pygofer with posterior
margin produced into dorsally directed spine,
spine not heavily sclerotized. Plate triangular,
lateral margin straight, apex rounded. Stylar
apex linear, preapical angle absent. Aedeagus
asymmetrical, left margin forming carina in
posterior aspect. Gonopore on right side near
base.

Type. — Male holotype, [Brazil] MS [Matto
Grosso du Sul], Campo Grande, l-X-1982,
W. KoUer collector. Type deposited in UPB.

Eri'MOLOGY. — Greek, haplo (simple) refers
to the color of this species.

Ileopeltus hastulus (DeLong &
Linnavuori), n. comb.

Figs. 27-32; Map 3

Chlorotettix hastulus DeLong & Linnavuori 1978:121-
122.

Diagnosis. — This species can be separated
from other members of this genus by the long
pygofer process and the aedeagus with a large
gonopore.

Length. — Male 5.0-5.2 mm, female un-
known.

Great Basin Naturalist Memoirs

No. 12

Figs. 19-22. Ileopeltus spinosus (DeLong): 19, pygofer, left lateral aspect; 20, right style, dorsal aspect; 21,
aedeagus, ventral aspect; 22, aedeagus and apex of connective, left lateral aspect.

Map 3. Distribution of //eo/«'/<u.s.s'pi»uwi<.s (squares), /. /iflp/i/s (circles), and/. /i«.sfi(/(/.v (triangles)

1988

CwiKLA: The Genus Ileopeltus

Figs. 23-26. Ileopeltus hapliis, n. sp.: 23, pygofer, left lateral aspect; 24, right style, dorsal aspect; 25, aedeagus,
ventral aspect; 26, aedeagus and apex of connective, left lateral aspect.

Color. — Ochraceous, without dark mark-
ings. Eye grey.

Male genitalia. — Pygofer with posterior
margin rounded in lateral aspect, long, dor-
sally directed process inserted on posteroven-
tral margin, processes crossing each other in
posterior aspect. Plate with lateral margin sin-
uate. Style posteriorly directed, preapical an-
gle rounded. Aedeagus with base expanded in
lateral aspect, small flange present on right
side in posterior aspect. Gonopore large on
right side at base of shaft.

Type. — Male holotype bearing the label
"Surumu, Roraima [Brazil], IX-1966, M. Al-
varenga and F. M. Oliveira Col." has been
examined from OSUC.

Distribution. — Known from the type lo-
cality (Roraima, Brazil) and a second male
specimen from Guarico, Venezuela, in
USNM. This species may be restricted to the
lowland tropics.

Ileopeltus blockeri, n. sp.
Figs. 33-37; Map 4

Diagnosis. — Ileopeltus blockeri is near 7.
hastulus and can be separated from it by the

pygofer processes inserted near the middle in
dorsal aspect.

Length. — Male 4.2-4.7 mm, female un-
known.

Color. — Ochraceous, without dark mark-
ings. Eye grey.

Male genitalia. — Pygofer with posterior
margin roundly produced, long acute process
inserted on ventral margin, near middle in
dorsal aspect, directed dorsally, crossing me-
dian line in dorsal aspect. Plate with lateral
margin insinuate, apex narrowly rounded.
Style linear, apex slightly curved laterally.
Aedeagus elongate compared to other mem-
bers of the genus.

Type. — Male holotype, Venezuela, Guar-
ico, lOkmeastofCalabozo, 18-III-1982, G. F.
Hevel and J. F. Hevel collectors. Type de-
posited in USNM.

Etymology. — This species is named in
honor of the noted leafliopper worker, H.
Derrick Blocker.

Ileopeltus clavatus , n. sp.

Figs. 38-42; Map 4

Diagnosis. — Ileopeltus clavatus can be

Great Basin Naturalist Memoirs

No. 12

Figs. 27-.32. Ueopeltus hastulus (DeLong & Linnavuori): 27, pygofer, left lateral aspect; 28, pygofer, posterior
aspect; 29, plates and valve, ventral aspect; 30, right style, dorsal aspect; 31, aedeagus, ventral aspect; 32, aedeagus and
apex of connective.

separated from other Ueopeltus species by the
truncate stylar apex that is shghtly constricted
subapically.

Length. — Male 4.2-4.4 mm, female 4.5
mm.

Color. — Ochraceous, without dark mark-
ings. Eye reddish brown.

Male genitalia. — Pygofer with posterior
margin truncate, stout process inserted on
posteroventral margin in lateral aspect, apex
of process crossing median line in dorsal as-
pect. Plate triangular, lateral margin straight,
fused to valve along anterior margin. Style
directed posteriorly, apex truncate, with
slight subapical constriction, preapical angle
absent. Aedeagus elongate, only slightly
asymmetrical.

Female seventh sternum. — Posterior
margin broadly convex, without excavations.

Type. — Male holotype, Brazil, RR [Ro-
raima], Boa Vista, 27-VII-1952, M. Alvarenga
collector. One female paratype, same data as
holotype. One male paratype, Venezuela,
Guarico, 10 km east of Calabozo, 18-III-1982,
G. F. HevelandJ. F. Hevel collectors. Holo-
type and female paratype deposited in UPB
and male paratype in USNM.

Etymology. — Latin, clavat (clubbed)
refers to the shape of the stylar apex.

Ueopeltus aberrans (Oshorn), n. comb.

Figs. 43-46; Map 4

Chlorotettix aherrans Osborn, 1923:72-73.
Doleranus cruzanus DeLong & Cwikla 1984:726. New
synonymy

Diagnosis. — Ueopeltus cruzanus is related
to /. cuneus and can be separated from it by
the narrow aedeagal shaft in ventral aspect.

Length. — Male 5.5-6.0 mm, female un-
known.

Color. — Ochraceous. Crown with faint
brown spot at apex of coronal sulcus, small
triangular patch approximately halfway be-
tween sulcus and eye. Pronotum ochraceous
with irregular, light brown spots on anterior
margin and six light orange bands running the
length of pronottuii. Forewing brownish sub-
h\'aline, dark brown coloration along commis-
sure on clavus and lew brown spots on middle
of wing.

Male genitalia. — Pygofer with posterior
margin truncate in lateral aspect, large, curved
spine on posteroventral margin, spine curves
medially then posteriorly, spines overlapping

1988

CwiKLA: The Genus Ileopeltus

Figs. 33-37. Ileopeltus blockeri, n. sp. ; 33, pygofer, left lateral aspect; 34, pygofer, dorsal aspect; 35, right style; 36,
aedeagus, ventral aspect; 37, aedeagus and apex of connective, left lateral aspect.

Map 4. Distribution of Ileopelttts blockeri (squdres), I. c/«ra/us (circles), and/. aZjerrans (triangles)

Great Basin Naturalist Memoirs

No. 12

Figs. 38-42. Ileopeltus clavatus , n. sp. : 38, pygofer, lateral aspect; 39, pygofer, dorsal aspect; 40, right style, dorsal
aspect; 41, aedeagus, ventral aspect; 42, aedeagus and apex of connective, left lateral aspect.

at middle. Plate triangular, slightly longer
than pygofer, lateral margin straight, not ap-
pearing fused to valve, two rows of setae
present, one row of macrosetae plus row of
small setaelike structures on lateral margin.
Stylar apex elongate, preapical angle absent.
Aedeagus U-shaped in lateral aspect, acute
apically. Gonopore basal, on right side of
shaft.

Type. — Male holotype bearing the labels
"Januaria, Minas Gerais, Brazil," "Dec 17,
1907" and "Carn. Mus. Ace. 3702" was exam-
ined from CMNH. The holotype oi criizanus
DeLong & Cwikla from Santa Cruz, Bolivia,
was examined from OSUC and found to match
the type of/, aherrans.

Distribution. — In addition to the type lo-
cality and Santa Cruz, Bolivia, two males from
Ceara and Sao Paulo, Brazil, were examined
from BMNH. The specimen from Sao Paulo
was collected from grass in a cloud forest.
Collecting dates included late February and
May.

Ileopeltus cuneus (DeLong &

Martinson), n. comb.

Figs. 47-50; Map 5

Chlorotettix cuneus DeLong & Martinson 1974:265.

Diagnosis. — Ileopeltus cuneus is close to /.

Cyclops and can be separated from it by the
asymmetrical aedeagus and the unique
pygofer process.

Length. — Male 4.8 mm, female unknown.

Color. — Ochraceous. Crown with median
triangular patch at apex of coronal sulcus, faint
brown spot halfway between sulcus and each
eye.

Male genitalia. — Pygofer produced into
long process, curved medially then dorsally.
Plate rounded apically, lateral margin slightly
convex. Stylar apex linear, spatulate, U-
shaped notch subapically on laterally surface
in dorsal aspect. Aedeagus slightly expanded
in posterior aspect, right side with two tooth-
like projections near base.

Type. — Male holotype bearing the labels
"Piracicaba, Sao Paulo, Brazil, 1-4-1966,"
"collr. C. A. Triplehorn," and "Blacklight
trap" was examined from OSUC.

Distribution. — Known only from the holo-
type.

Ileopeltus Cyclops (Linnawiori), n. comb.

' Figs. 51-,55; Map 5

Doleranus cyclops Linnavuori 1959:275-276.

Diagnosis. — This unique species can be
separated from other Ileopeltus species by the
median spot on the crown and the unique
stylar apex.

1988

CwiKLA: The Genus Ileopeltus

Figs. 43-46. Ileopeltus aberrans (Osborn): 43, pygofer, left lateral aspect; 44, right style, dorsal aspect; 45, aedeagus,
ventral aspect; 46, aedeagus and apex of connective, left lateral aspect.

Length. — Male 4.9 mm, female unknown.

Color. — Ochraceous. Crown with median
triangular brown patch. Eye gray.

Male genitalia. — Pygofer with posterior
margin produced into acute process posteri-
orly directed then curving medially, sclero-
tized. Plate short, rounded apically. Stylar
apex acute, distal third abruptly curved later-
ally, preapical angle absent. Aedeagus only
slightly asymmetrical, shaft narrow in poste-
rior aspect.

Type. — Male holotype bearing the labels
"Loreto, Missiones [Lapsus calmi for Mi-
siones? Possibly from Corrientes], ARC [Ar-
gentina] XII-4-1931, A.A. Ogoblin" and
"collected at light" has been examined from
USNM.

Distribution. — Known only from the holo-
type.

Cladistic Relationships among the Species

Biological classifications should be based on
hypothesized relationships estimated by cla-
distic (or phylogenetic) analyses (Hennig
1966). Such analyses are dependent upon the
determination of plesiomorphic and apomor-
phic character states. Hypotheses of character

polarity are accomplished by comparing the
taxa under question with their outgroup.
Ileopeltus is thought to be closely allied to
Chlorotettix by nature of both having the
crown completely microsculptured. Because
of this single synapomorphy, Chlorotettix is
considered the sister-group to Ileopeltus .

Twelve binary characters were chosen for
the cladistic analysis and are summarized in
Table 1. The resulting data matrix (Table 2)
was analyzed by means of phylogenetic analy-
sis using the parsimony (PAUP) program de-
veloped by Swofford (1985).

The paucity of characters for Ileopeltus spe-
cies, I believe, is the result of a generalized
reduction in complexity of characters. This
becomes immediately obvious when the male
genital structures are examined, e.g., the
aedeagus without processes. These reduced
characters most assuredly obscure the inter-
relationships among the species and possibly
at higher levels. Even with this reduction,
most of the characters used in this analysis
were genitalic and are discussed below.

The hypothesized cladogram is presented
in Figure 56. The most primitive lineage con-
tains the species /. tethys and /. nanocanthus ,

Great Basin Naturalist Memoirs

No. 12

Figs. 47-50. lleopettus citneus (DeLong & Martinson): 47, pygofer with anterior portion damaged, left lateral aspect;
48, right style, dorsal aspect; 49, aedeagus, ventral aspect; 50, aedeagus and ape.x of connective, right lateral aspect.

Map 5. Distribution oi Ileopeltus cuiwtis (sciuarcs) and /. ctjclops (circk

1988

CwiKLA: The Genus Ileopeltus

Figs. 51-55. Ileopeltus cyclops (Linnavuori): 51, pygofer, left lateral aspect; 52, right style, dorsal aspect; 53,
aedeagus, ventral aspect; 54, aedeagus and apex of connective, left lateral aspect; 55, head, pronotum, and scutellum,
dorsal aspect.

Fig. 56. Hypothesized cladogram for the genus Ileopeltus. Numbers refer to characters listed in Table 1.

Great Basin Naturalist Memoirs

No. 12

Fig. 57. Ileopeltus tethys (Van Duzee), aedeagus, ventral aspect. Fig. 58. /. nanocanthus, n. sp. , aedeagus, ventral
aspect.

which share the synapomorphy of the concave
plate and the style with the preapical angle
absent. Both characters are homoplastic and
are found throughout the cladogram. The re-
maining species have a distinct process.

Of those Ileopeltus species with prominent
pygoferal processes, two major lineages are
hypothesized. One contains those individuals
with stout pygoferal processes; the other is
defined by long, heavily sclerotized pro-
cesses.

The lineage with stout pygoferal processes
contains four species. Ileopeltus dorsalus has
a median black patch on the crown, while the
remaining three species have the preapical
angle of the style absent, a homoplastic char-
acter found throughout the cladogram.
Ileopeltus haplus and /. spinosus have their
stylar apex truncate, another homoplastic
character.

The lineage that is defined by the long
sclerotized process is also supported by the
processes curved medially, a character which
is later reversed. The lineage itself is repre-

sented by an unresolved trichotomy contain-
ing/, hastulus, I. blockeri, and the remaining
Ileopeltus species. Ileopeltus hastulus is de-
fined by the autapomorphy of a large gono-
pore. The remaining four species have the
preapical angle of the style absent. As men-
tioned before, this is a homoplastic character.
Ileopeltus aberrans, I. cy clops, and I.
cuneus have a median black patch on their
crown and the lateral margin of their plate
concave. Both characters are homoplastic.
Only autapomorphic characters define these
species.

Acknowledgments

I thank the following individuals and their
associated institutions for allowing me to ex-
amine specimens under their care: M. D.
Webb, Department of Entomology, British
Museum (Natural History), London (BMNH);
C. W. Young, Section of Entomology, Car-
negie Museum of Natural History, Pittsburgh
(CMNH); C. A. Triplehorn, Department of

1988

CwiKLA: The Genus Ileopeltus

Table 1. Characters, presumed plesiomorphic and
apomorphic states for Ileopeltus species, and numbers
that refer to character distribution presented in Table 2
and Fig. 56.

Fig. 59. Ileopeltus tethys (Van Duzee), aedeagus, right
lateral aspect. Fig. 60. Ileopeltus nanocanthus, n. sp.,
aedeagus, right lateral aspect.

Entomology, Ohio State University, Colum-
bus (OSUC); the late Francisco Jose Fer-
nandes-Yepez, Facultad de Agronomia, Uni-
versidad Central de Venezuela, Maracay
(UCV); Paul H. Freytag, Department of
Entomology, University of Kentucky, Lex-
ington (UKC); K. M. Rocha Zanol, Universi-
dade Federal do Parana, Curitiba, Brazil
(UPB); J. P. Kramer and R. Froeschner, De-
partment of Entomology, Smithsonian In-
stitution, Washington (USNM); H. Wolda,
Smithsonian Tropical Research Institute, Bal-
boa, Republic of Panama (HWC). H. Derrick
Blocker (Kansas State University), P. H.
Freytag (UKC), Sam Stribling (USNM), and
G. M. Chippendale (University of Missouri,
Columbia) kindly criticized an earlier draft of
this manuscript. Niki K. J. Miller (Ohio State
University) skillfully executed the habitus
drawing. Special thanks are extended to Paul
W. Oman, whose works on leafhopper sys-
tematics have proved to be most inspiring.

Plesio-

Apo-

Character

morphic

Coloration

unmarked

crown marked
with median
triangular
patch

Pygoferal processes

absent

present

Pygoferal processes

elongate

yes

Pygoferal processes

stout

yes

Pygoferal processes long

and heavily sclerotized

yes

Pygoferal processes
strongly curved

medially

yes

Plates fused to valve

yes

Plates with lateral

margin

straight

concave

Style with preapical

angle

present

absent

10.

Stylar apex

stout

narrow

11.

Stylar apex

rounded

truncate

12.

Gonopore

small

large

Literature Cited

Ball. E G 1936. Some new leafhoppers related to Tham-
notettix. J. Washington Acad. Sci. 26: 430-434.

Caldwell, J S , and L F Martorell 1950. Review of
the Auchenorynchous Homoptera of Puerto Rico.
Part 1. Cicadellidae. Puerto Rico Univ. J. Agric.
34: 1-132.

CwiKLA, P. S. 1988. A systematic study of the leafhopper
genus Chlorotettix (Homoptera: Cicadellidae).
Unpublished dissertation, Ohio State University,
Columbus.

CwiKLA, P S , AND P. H Freytag 1983. External mor-
phology of Xestocephalus subtessellatus (Ho-
moptera: Cicadellidae: Xestocephalinae). Ann.
Entomol. Soc. Amer. 76: 641-652.

DeLong. D M. 1923. New species. Pages 257-270 in
G. N. Wolcott, Insectae Portoricensis. Puerto
Rico Univ. J. Agric. Vol. 7.

1945. The genus Chlorotettix (Homoptera: Ci-
cadellidae) in Mexico. Lloydia8: 1-30.

DeLong, DM, and P S. Cwikla 1984. New Neotropi-
cal Deltocephalinae (Homoptera: Cicadellidae). J.
Kansas Entomol. Soc. 57: 725-728.

DeLong, D M , andC Martinson 1974. New species of
Chlorotettix from Argentina, Bolivia, Brazil, and
Mexico. J. Kansas Entomol. Soc. 47: 261-267.

DeLong, D M , and R E Linnavuorl 1978. Studies on
Neotropical leafhoppers (Homoptera: Cicadelli-
dae). Entomol. Scand. 9: 111-123.

Hennig, W 1966. Phylogenetic systematics. University
of Illinois, Urbana.

Linnavuorl R E 1959. Revision of the Neotropical Del-
tocephalinae and some related subfamilies (Ho-
moptera: Cicadellidae). Ann. Zool. Soc. Vanamo

Great Basin Naturalist Memoirs

No. 12

Table 2. Data matrix used in constructing a cladogram for Ileopeltus. Characters number refers to information in
Table 1: 0, plesiomorphic; 1, apomorphic.

Character

Chlorotettix

1. cuneus

I. Cyclops

I. aherrans

1. blockeri

I. hastulus

I. tethijs

I. nanocanthus

I. haplus

I. dorsahis

1. clavatus

I. sphwsus

I. ventriculus

20: 1-370.

OSBORN. H 1923. Neotropical Homoptera of the
Carnegie Museum. Part 2. Records and descrip-
tions of five new genera and sixty-five new species
of the subfamily Jassinae. Carnegie Museum Ann.
1.5: 27-79.

1935. Insects of Puerto Rico and the Virgin Is-
lands. Homoptera (excepting the Sternorhynchi).
Puerto Rico and Virgin Islands Sci. Surv. 14:
111-260.

Oman, P W 1949. The Nearctic leaflioppers (Homop-
tera: Cicadellidae); a generic classification and
check list. Mem. Entomol. Soc. Washington No.
3. 2.53 pp.

SwoFFORD, D L 1985. PAUP, phylogenetic analysis us-
ing parsimony. Users manual. Illinois Natural
History Survey, Champaign.

Van Duzee. E P 1907. Notes on Jamaican Hemiptera: a
report on a collection of Hemiptera made on the
island of Jamaica in the spring of 1906. Buffalo Soc.
Nat. Sci. Bull. 8: .3-79.

WOLCOTT. G N 1923. Insectae Portoricensis. A prelimi-
nary annotated check-list of the insects of Puerto
Rico, with descriptions of new species. Puerto
Rico Univ. J. Agric. 7: 1-313.

1936. Insectae Borinquenses. A revised annotated

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Univ. J. Agric. 20: 1-600.

SOME ASPECTS OF THE BIOLOGY, MORPHOLOGY, AND EVOLUTION OF
LEAFHOPPERS (HOMOPTERA: CICADELLOIDEA AND MEMBRACOIDEA)

J. W. Evans'

Abstract — This article summarizes some observations of a varied nature on the biology, morphology, and evolution
of the Cicadelloidea (Cicadellidae, Hylicidae, Eurymelidae) and Membracoidea(Membracidae, Aetalionidae, Biturri-
tidae, Nicomiidae). These observations, made over a period of more than half a century, have previously been recorded
at different times, but lie buried in the literature. It is hoped that their interest will justify repetition and draw attention
to some promising lines of research.

Biology
Food Plant Associations

As Southwood (1961) has pointed out, in-
sects have a particularly close association with
plants belonging to the predominant flora of
the time. Thus, most Australian cicadelloids
feed on eucalypts and numerous membracids
on acacias. Of particular interest are relict
insects that feed on representatives of ancient
floras. Examples are Koebelia californica
Baker on conifers in California and Cornutipo
spp. (Eurymelidae) on representatives of the
Proteaceae in Australia. The Cephalelini, a
tribe of the Ulopinae which have an Antarctic
distribution, provide another example, for
their feeding seems to be restricted to an an-
cient family of plants, the Restionaceae, in
Australia, South Africa, and New Zealand.

Changes of food plants have recently oc-
curred in the Tartessinae (Cicadellidae).
These are primarily eucalypt feeders, but
some have become adapted to living in tropi-
cal rain forests and have taken to feeding on
other kinds of trees (F. Evans 1981).

Sound Production and Courtship

Ossiannilsson, in his notable study of the
"songs" of Homoptera, was the first to draw
attention to the presence of tymbals in insects
of both sexes belonging to the Cicadelloidea,
Cercopoidea, and Cicadoidea (Ossiannilsson
1949). Previously, I had noted the presence of
structures that I described as "resembling
tymbals" on the first abdominal segment of
Darthula hardwickii Gray and Aetalion retic-

ulatum Linnaeus (Evans 1946b). In his discus-
sion of the function of the songs of various
Auchenorrhyncha, Ossiannilsson described
some as being "calls of courtship." Subse-
quently, I noted the presence of well-devel-
oped tymbals in nymphs belonging to every
instar but the first in both sexes of A. reticida-
tiim (Evans 1957). As, presumably, nymphal
song cannot have a sexual significance, it may
serve an assembling purpose since aetalionids
are gregarious insects. However, amongst the
Eurymelidae, which are likewise gregarious,
I have been unable to find any trace of tym-
bals. Nevertheless, prior to mating, eu-
rymelids indulge in a prolonged courtship
(Evans 1931).

Tymbal sound production occurs among
representatives of all the superfamilies of the
Auchenorrhyncha, and even in the
Cicadoidea it is not confined to one sex since
females of Tettigarcta can "sing." Thus, it
must have been a feature acquired in early
Mesozoic, if not Palaeozoic, times (Evans
1941). Moreover, it would seem that during
the early Mesozoic some Auchenorrhyncha
were capable of stridulation. This is supposed
because some Upper Triassic Homoptera
(e.g. , Eoscartoides bryani Evans) have stridu-
latory areas on the proximal costal area of their
tegmina (Evans 1961).

Oviposition

When ovipositing, the majority of leafhop-
pers insert their eggs into plant tissue, either
singly or in batches. In the latter case, the
eggs are covered with secretions that harden

'Australian Museum, Sydney, N.S.W., Australia. Present address; 47 Bundarra Road, Bellevue Hill, Sydney 2023, Australia.

Great Basin Naturalist Memoirs

No. 12

on exposure to the air. The eggs of aetahonids
and of some bitturitids and membracids are
contained in oothecae situated on the surface
of the plants. Because the AetaHonidae are an
ancient group of insects, it might be thought
that such a method of oviposition predated
egg insertion in plant tissue. However, this is
improbable, as the last-named method is
shared with insects belonging to the super-
families of the Auchenorrhyncha; and eggs,
whether laid inside plants or on their surface,
are equally subject to heavy parasitism.

Gregariousness and Ant Attendance

While all eurymelids and aetalionids, many
membracids, and possibly all bitturitids are
gregarious, so far as I am aware, no cicadellids
have this behavior pattern. Since female
membracids sometimes remain with their
eggs after oviposition, some authors, such as
Haviland (1925), have credited them with ma-
ternal care. Eurymelids of both sexes have
been recorded as remaining with their eggs
for a period. It is possible that the gregarious
habit in this family may be associated with the
fact that their nymphs, unlike those of other
leafhoppers, lack the ability to jump.

It would seem that all gregarious leafliop-
pers are ant attended, but the Pogonoscopini,
a tribe of the Eurymelidae, are the only ones
that have developed the characteristics of true
myrmecophiles; i.e., they have unusually
long legs and small eyes and live in the nests of
ants (Evans 1966).

Morphology
Color and Size

Previously, 1 have suggested that early
leafhoppers were brown, as, for instance, is
Darthula hardwickii , and that green was the
first alternative color to be acquired (Evans
1966). Later evolutionary developments have
given rise to insects that may be predomi-
nantly black and have, or lack, yellow, red, or
occasionally bluish markings. Some Cicadelli-
dae from Madagascar have striking color pat-
tern differences (Evans 1953). In others, spe-
cies occur that have identical male genitalia
and comprise populations with differing, but
stable, color patterns (e.g., Eunjmela fenes-
trata Le Peletier & Serville) (Evans 1933).
Then, some species have individuals with a
very variable color pattern (e.g., Euryme-

loides punctata [Signoret]) (Evans 1966).

The largest known leaffiopper, the aetal-
ionid Darthula hardwickii Gray, has a length
of 28 mm, of which the apical 12 mm in both
sexes consists of the prolonged ninth abdomi-
nal segment. The smallest leafhoppers are
comprised in the Typhlocybinae (Cicadelli-
dae), some species of which are no more than
2 mm long. Evolutionary development seems
to be frequently accompanied by increase in
size. Thus, for example, in the Eurymelidae,
the largest species are seemingly the most
recently evolved (e.g., Eunjmelops generosa
[Stal]).

The two sexes of Cicadelloidea usually re-
semble each other in coloration and size. An
exception in respect to color is provided by
the sole representative of the Tartessinae to
occur in New Caledonia. This insect, Calo-
tartessus stalii (Signoret), has males that are
largely black and females that are predomi-
nantly brown. In regard to size, the Stenoco-
tini (Ledrinae) have males that are consider-
ably smaller than females.

The Head

The most puzzling feature of the heads of
Hemiptera is associated with the origin of
their feeding apparatus. Insects in some other
orders feed by suction, but with all these the
nature of the transition from mandibulate to
haustellate mouthparts can be readily under-
stood.

Previously, 1 have suggested that, though
the ocelli are on the crown in representatives
of the relict subfamily, the Ulopinae, this con-
dition is a secondary one. If this is correct,
even though insects with ventral ocelli do not
seem to be disadvantaged, such a change of
position would seem to be an adaptive one.

Other primitive features of leaffiopper
heads are the presence of a sensory pit on the
maxillary plates, which may possibly be
derived from the maxillary palp (Evans 1973),
and the complete separation of the maxillary
plates from the genae.

Male Genitalia

The male genitalia of insects are more sub-
ject to change than any other parts of their
bodies. For this reason, they are extensively
used for species recognition. The nature and
the extent of observable differences between
the aedeagi of closely related insects vary

1988

EvANS; Leafhopper Biology, Morphology, and Evolution

widely and range from slight and constant to
considerable and variable ones. The subgeni-
tal plates and accessory processes are also sub-
ject to considerable change of shape.

While male genitalia differences are useful
for species separation and genus recognition,
they are sometimes also helpful for family
recognition purposes. Thus, for example,
while within the Cicadellidae the aedeagus
invariably arises from a basal connective situ-
ated between the paired parameres, in the
Eurymelidae it lacks any association with the
basal plate and the parameres.

Inasmuch as simple male genitalia serve the
same purpose as complex ones and pre-
sumably in an equally satisfactory manner,
and the structure of female genitalia is con-
stant, it is difficult to understand the nature of
the advantages conferred by increasingly
complex male genitalia unless female insects
avoid mating with males that have the wrong
"key. "

Adaptive Characters

In addition to the ocelli noted above, other
adaptive characters among leaflioppers are to
be found in the Stenocotini, an endemic Aus-
tralian subfamily of the Ledrinae, and in the
Eurymelidae. Stenocotids, which live under
bark of eucalypts, are flattened insects, and
their nymphs are paper thin. While most eu-
rymelids are wedge-shaped, one species,
Platyeurymela semifascia (Walker), which,
like stenocotids, lives under bark, is oval in
shape and convex.

Alary dimorphism and polymorphism are of
widespread occurrence in the Heteroptera,
they are rare in the Homoptera-Auchenor-
rhyncha, and in the Cicadellidae they would
seem to be particularly associated with insects
living at high altitudes or in a marsh environ-
ment. Thus, Monteithia spp. (Monteithiini,
Ulopinae), which live at high altitudes in New
Guinea, and of which fully winged insects
have not been recorded, have males that dif-
fer from females in the extent to which their
wings are foreshortened (Evans 1968). Then,
in Cephalelini, which inhabit both alpine and
low-level marsh environments, both sexes oc-
cur in a flightless and a fully winged condition.
Some other leaflioppers with both brachyp-
terous and fully winged individuals are
Taslopa montana Evans (Ulopini, Ulopinae),
Chiasmus varicolor (Kirkaldy) (Deltocephali-

nae), and Euacanthella sp. (Euacanthellinae).

The best-known examples of the acquisition
of morphological features among leaflioppers,
which are seemingly of nonadaptive signifi-
cance, are to be found among the Mem-
bracidae. This phenomenon has been dis-
cussed by many authors (e.g., Haviland 1925)
who observed these insects in the territory
then known as British Guiana. She pointed
out that the early stages of exaggerated prono-
tal development in insects, which, in their
extreme form, were either cryptic or mimetic,
cannot have conferred any protective advan-
tage, and yet the insects survived. Moreover,
the pronota of many membracids seem to lack
any degree of protective significance.

Above I have suggested that the extreme
differences found among leaflioppers belong-
ing to the genus Colloborrhis Germar in
Madagascar may have been initiated by
"explosive speciation." I have also formerly
suggested that the same phenomenon may
have occurred among Membracidae isolated
in the Neotropical Region during Tertiary
times (Evans 1959). This is because, though
doubtless enlarged pronota were a mem-
bracid characteristic before this isolation took
place, it is in South America only that the
evolution of such a range of bizarre forms took
place.

The development in leaflioppers of bizarre
structures, which apparently lack adaptive
significance, is not confined to changes in the
shape of the prothorax. These occur also in the
heads of leaflioppers, as, for example, in those
of Cornutipo tricornis Evans, Listrophora
evansi Evans, and Wolfella caternaulti
Spinola (all illustrated in Evans 1975a).

Evolution
Fossils and the Classification of Recent Forms

An abundance of wings of Homoptera has
been found in Permian and Triassic strata in
both the northern and southern hemispheres.
These seem to provide evidence that all the
existing superfamilies of the Auchenorrhyn-
clia were already differentiated by Triassic
and possibly Permian times. If this suggestion
is valid, it provides an example of unusual
evolutionary stability (Evans 1964).

In the forewings, or tegmina, of some
Upper Permian Homoptera (e.g., Homa-
loscytina plana Tillyard) six veins support

Great Basin Naturalist Memoirs

No. 12

their apices. These are as follows: Rib, Rs,
Ml, M2, M3, and M4. In some recent Ci-
cadelloidea, only four veins serve the same
purpose, and it is of interest to note that the
identity of these is different in each of the
comprised families. Thus, in the Hylicidae,
they are Rib, Rs, Ml + 2, and M3+4 + CuA;
in the Eurymelidae, Rl, Ml + 2, M3+4,
and CuA; and in the Cicadellidae, Rib,
Rs + Ml+2, M3 + 4, and CuA. This circum-
stance would seem to provide evidence of the
separate, direct derivation of insects com-
prised in the leafhopper families from a com-
mon ancestral stock (Evans 1949, 1964). The
recently found fossil insect placed in a new
Homoptera family, the Jascopidae, belongs in
my opinion to the Cicadellidae (Evans 1972).

Parallel Evolution

The most striking example of parallel evolu-
tion in the family Cicadellidae is provided by
the resemblance between insects comprised
in the Cephalelini (Ulopinae) and the Para-
dorydiini (Hecalinae). Leafhoppers in both
tribes have species with short, triangular
heads and others with long, narrow ones, the
complete insect being seedlike in appearance.
While the Paradorydiini, unlike the Cephale-
lini, do not feed on the rushlike Restionaceae,
some have been recorded from "rushes ' grow-
ing in marshes. The Ulopinae and the Hecali-
nae are not closely related and probably were
differentiated during different geological pe-
riods, the former being of Mesozoic and the
latter possibly Tertiary origin.

Explosive Speciation

Above I have mentioned an instance of sup-
posed "explosive speciation that has occurred
in Madagascar with leafhoppers belonging to
the genus Coloborrhis Germar (Ulopini, Ulo-
pinae). The type species of the genus, C. cor-
ticina Germar, has an extensive distribution
in Africa from where no other species has
been recorded. It is established also in Mada-
gascar, from where no less than 16 other spe-
cies, which could equally well be regarded as
belonging to separate genera, have been de-
scribed. These differ strikingly from one an-
other in characters of the head and thorax.
Thus, the head may be rounded anteriorly, or
narrowly, or broadly spatulate, and it may
have differently shaped prominences on the
crown. Then, the pronotum may have a pair of

lateral prominences of varying size, or be lat-
erally humped, and it may be widest either
anteriorly or posteriorly, while the scutellum
may be of normal size or raised into a large,
vertical crest. The tegmina may have basic
cicadellid, or reticulate, venation. One spe-
cies (C. rugosa Evans) has a characteristic
shared only with the tegmina of Permian and
Triassic Homoptera. This is the proximal
arching of vein CuA so that it makes contact
with vein M. Finally, the hind tibae may be
narrowly rectangular in section, or broadly
spatulate. It is of interest to note that some of
the characteristics listed above are shared
with the Ledrinae, while others are of a mem-
bracidlike nature (Evans 1953, 1959).

Supposed Sympatric Speciation

During 1959 a symposium was held in Mel-
bourne to celebrate the centenary of the Royal
Society of Victoria. Its proceedings were later
published under the title "Evolution of living
organisms." At the end of the paper I con-
tributed to this symposium (Evans 1962), I
said:

The concept of sympatric speciation is at the present
time ahnost universally discredited and any mention of it,
even as a possibility, might seem to have no place in a
contribution purporting to be scientific. This is especially
so when, as in the case of the present instance, a sugges-
tion is made that is unsupported by experimental evi-
dence. Nevertheless, in my opinion, it is possible that
sympatric speciation may take place within a particular
group of leaflioppers, the Typhlocybinae, and the reason
for this opinion is because amongst these insects disconti-
nuities between populations of an ecological and ethologi-
cal nature would seem to be more readily capable of
achievement than isolation of a geographical nature.

Typhlocybids, which are of world-wide distribution,
are particularly well represented in the Holarctic Region.
They range in length from 2-4 mm. This group comprises
some hundred genera, several of which contain many
hundreds of distinct species. These leaflioppers differ
greatly in the extent to which they are restricted in their
feeding requirements. Some feed, mate and breed only
on a single species of plant or on a limited range ol related
plants. Others have wider feeding habits. The mere fact
that related species have different ranges of food plants
implies an ability, on occasion, to become accommodated
to a new diet.

Mention has already been made of the songs of Ho-
moptera and it may be of significance that typhlocN bids
have unusually large apodemes for the support of their
tymbal muscles and hence, presumably, are particularK'
vocal. It is accordingly suggested that an isolating factor
which may have enabled the evolution of large numbers
of sympatric species might be the s\nchronization of the
ac(iuisition of a new call note with that of a new food plant.

The reason it has been suggested that it is improbable

1988

EVANS: LEAFHOPPER BiOLOCJY , MOKPMOLOGY, AND EVOLUTION

that geographical isohition has been tlie principal factor
enabhng prohfic speciation to take place among typhlocv -
bids is because their small size and swarming habits make
them particularly liable to transport in the upper air and
hence to rapid and wide dispersal.

My views on the above matter have not
changed during the 25 years that have elapsed
since I first expressed them.

Some Characteristics of
Island and Montane Faunas

When the gene flow of insects is restricted
following the isolation of small initial popula-
tions, the process of evolution is often acceler-
ated. Mention has already been made of de-
velopments that have occurred on the island
of Madagascar following the presumed isola-
tion of a population of the ulopid Coloborrhis
corticina . The occurrence of aberrant leaf-
hoppers is not confined to large islands. Thus,
on Juan Fernandez Island there occurs possi-
bly the most grotesque of all leafhoppers,
Evansiola kiischeli (China). The head of this
species is twice as wide as long, and, when
observed from above, projects laterally con-
siderably beyond the sides of the rest of the
body (Evans 1975a).

Montane faunas in Australia and New
Guinea comprise survivors from a time when
prevailing climates were cooler than at pres-
ent and from immigrant species belonging to
later evolved groups. Examples belonging to
the first category are provided by Taslopa
montana Evans (Ulopinae, Ulopini), which
has been recorded only from high altitudes in
Tasmania and southeastern Australia, and
Monteithia spp., which inhabit high-level en-
vironments in New Guinea (Evans 1966,
1968). A leafhopper in the second category is
Atistroagalloides rosea Evans (Austroagal-
loidinae) (Evans 1966).

Relict Characteristics

The relict characteristics of leafhoppers can
be recognized by making a comparison of
those features in which presumed recently
evolved forms differ from those of presumed
earlier origin.

The head. — Below are given particulars of
how recent Deltocephalinae differ in head
structure from the ancient aetalionid,
Darthula hardwickii. Not only are the Del-
tocephalinae probably the most recently
evolved of all cicadellids, but they are also the

most widely distributed and contain the
largest number of species. The characteristics
of representatives of this subfamily, given be-
low, precede those of D. hardwickii, which
are given in parentheses. Wide maxillary
plates with sensory pits, continuous with the
genae (narrow maxillary plates with sensory
pits, separated from the genae by a subgenal
suture); hind margin of lora widely separated
from the antennal bases (hind margins of lora
adjacent to the antennal bases); frontoclypeus
lacking any trace of epistomal suture and ex-
tending posteriorly as far as the hind margin of
the face (frons separated from the postclypeus
by an epistomal suture anteriorly and the
arms of the epicranial suture posteriorly);
ocelli situated near the sides of the fronto-
clypeus and close to, or on, the margin of the
head separating the face from the crown
(ocelli on the face of the head close to the sides
of the epicranial suture and at a considerable
distance from the hind margin of the face)
(Evans 1975a).

The thorax. — Some supposed relict char-
acters are the small pronotal paranota, such as
occur in the Myerslopiini (Ulopinae), and ex-
tensions of the costal region of the tegminal
pads of the nymphs of some ulopids, such as
those of Coloborrhis corticina and also a few
macropsids (Evans 1968). These paranota and
the mesothorasic costal expansions are re-
garded as relict features because the former
were possessed by many Palaeozoic insects
and the latter occur also in the Psylloidea, and
in some Cicadelloidea and Membracoidea,
which, for other reasons, are regarded as
relict forms.

The abdomen. — Among relict characters is
the retention of tymbals in the nymphs as well
as in the adults of some aetalionids, and the
presence in the male genitalia of bisegmented
subgenital plates. I have suggested that the
latter, which occur in some Ulopinae,
Macropsinae, and Agallinae, represent the
gonocoxites and the gonostyli of the ninth ab-
dominal segment (Evans 1975).

Conclusions

The observations recorded in this article
serve to emphasize that the factors responsi-
ble for evolutionary change are of varied na-
ture. Some, involving major changes of struc-
tural organization, present puzzling features.

Great Basin Naturalist Memoirs

No. 12

Others suggest that evolutionary develop-
ments need not always be of an adaptive na-
ture, that isolation of populations need not
always be of a geographical nature, and that
the development of bizarre forms is particu-
larly associated with isolation of populations
on islands.

Literature Cited

Evans, F. 1981. The Tartessinae of Australia and New

Guinea (Homoptera, Cicadellidae). Pacificlnsects

23: 112-188.
Evans, J. W 1931. Notes on the biology and morphology

of the Eurymelinae. Proc. Linn. Soc. N.S.W. 56:

210-226.
1933. A revision of the Eurymelini (Homoptera,

Bythoscopidae). Trans. R. Soc. S. Australia 57:

73-90.
1941. The morphology of Tettigarcta toinentosa

White. Pap. Roy. Soc. Tasmania 1940; 35-49.
1946a. A natural classification of lealhoppers Qas-

soidea, Homoptera) Pt. I. Trans. R. Soc. London

96: 47-60.
1946b. A natural classification of lealhoppers Pt.

II. Trans. R. Soc. London 97: 39-54.
1953. Les Cicadellidae de Madagascar. Mem.

Inst. Sci. Madagascar E. 4: 87-137.
1957. Some aspects of the morphology and inter-
relationships of extinct and recent Homoptera.

Trans. R. Entomol. Soc. London 169: 275-294.
1959. Quelques nouveaux Cicadellides de Mada-

gascar. Mem. Inst. Sci. Madagascar E. 11: 481-
507.

1961. Some Upper Triassic Heniiptera from

Queensland. Mem. Queensland Mus. 14: 13-23.

1962. Evolution in the Homoptera. Pages 250-

259 in The evolution of living organisms. Mel-
bourne University Press.

1964. The periods of origin and diversification of

the superfamilies of the Auchenorrhyncha as de-
termined by a study of the wings of Palaeozoic and
Mesozoic fossils. Proc. Linn. Soc. London 175:
171-181.

1966. The lealhoppers and froghoppers of Aus-
tralia and New Zealand. Mem. Australian Mus.
12: 1-347.

1968. Some relict New Guinea lealhoppers and

their significance in relation to the comparative
morphology of the head and thorax of the Ho-
moptera-Auchenorrhvncha. Pacific Insects 10:
215-229.

1972. Some remarks on the family Jascopidae (Ho-
moptera, Auchenorrhyncha). Psyche 79: 120-121.

1973. The maxillary plate of Homoptera- Auchen-
orrhyncha. J. Entomol. 48: 43-47.

197.5a. The external features of the heads of

lealhoppers. Rec. Australian Mus. 29: 407-439.

1975b. The structure, function and origins of the

subgenital plates of leafhoppers. J. Australian En-
tomol. Soc. 14: 77-80.

Haviland, M D 1925. The Membracidae of Kartabo,
British Guiana. Zoologica 6: 239-290.

OssiANNlLSSON, F 1949. Insect drummers. Opusc. Ento-
mol. Suppl. 10: 1-142.

SouTHWOOD, T R E 1961. The numbers of species of
insects associated with various trees. J. Anim.
Ecol. 30: 1-8.

REVISION OF THE GENUS CALLISCARTA (HOMOPTERA:
CICADELLIDAE: NEOBALINAE)'

Paul H. Freytag^

Abstract. — A taxonomic revision of the Neotropical genus Calliscarta Stal is presented. Most of the 19 species
treated are described and illustrated, including 10 that are new to science. The new species include: ahrupta (Bolivia),
acuta (Peru, Venezuela), delicata (Bolivia, Peru), elon^ata (Ecuador, Peru), expansa (Bolivia, Ecuador, Peru), lora
(Peru), iimrginata (Honduras), inexicana (Mexico), ornata (French Guiana, Venezuela), riig,osa (Venezuela). Three
cases of new synonymy are proposed; boliviana (Osborn) (- tiixfia Kramer), /a.sctafa (Oshorn) [= brunnea (Osborn)],
decora (Fabricius) [= magnifica (Osborn)].

The genus Calliscarta as reviewed by Lin-
navuori (1959) included nine species. Two
species and the genus Idiotettix Osborn were
placed in synonymy. Since then Linnavuori
and Heller (1961) added one species, and
Kramer (1963) added two species. In this
study nine of these species are recognized as
valid, three are placed in synonymy, and 10
new species are added. The limited number
of specimens in collections of this genus make
it a very difficult genus with which to deal. No
division of this genus is made at this time even
though some species groups appear to repre-
sent separate genera, such as the fasciata
group. Much more material will be needed,
especially from Central America, to know how
to properly place many species. The problem
of having only one sex of the majority of the
species also makes it very difficult to under-
stand relationships. For these reasons a phy-
logenetic study must wait until the species are
better known.

Calliscarta Stal

Calliscarta Stal 1869: 82 (type-species Cicada decora
Fabricius); Dallas 1870: 497; Van Duzee 1890: 35;
Van Duzee 1890: 79; Schulze, Kukenthal, and
Heider 1927: 504; Neave 1939: 537; Evans 1947:
230; Linnavuori 1959: 27; Linnavuori and Heller
1961:3;Metcalfl967:946.

Idiotettix Osborn 1929: 465 [type-species Thamnotettix
magnificus Osborn { = Cicada decora Fabricius)]
Oman 1938: 355; Neave 1939: 763; Metcalf 1944
161; Evans 1947: 192; Evans 1951: 9; Nast 1952: 1
Maldonado Capriles 1954: 247; Metcalf 1966: 225.

Large, robust, wedge shaped. Head wider
than pronotum; crown wide, same length near

eyes as median, coronal suture short; ocelli on
the margin about twice their width from eyes,
face elongate; postclypeus convex and long,
sutures along each side starting near ocelli,
with antennal ledges from postclypeus to eye.
Pronotum convex, lateral margins short, pos-
terior margin slightly but evenly emarginate.
Forewing truncate, appendix well developed,
with three subapical cells. Femur of hind leg
with spinulation 2-2-1. Male genitalia large
and mostly sclerotized. Anal tube long, cylin-
drical, heavily sclerotized. Pygofer always
with a membranous ventral process that ex-
tends into the genital capsule, or with a long
ventral process.

This genus has been previously placed in
the subfamily Idiocerinae on the basis of the
overall shape of the body and the large ap-
pendix. It has also been placed in the subfam-
ily Deltocephalinae, as the ocelli are on the
margin of the head near the eyes. These and
other characters, plus the fact that the major-
ity of the species are brightly colored and
larger than those found in the above-men-
tioned subfamilies, caused Linnavuori (1959)
to place this genus in the subfamily Neobali-

Key to Known Males

1. Aedeagus with extra dorsal apodemal process
(Figs. 56, 57); pygofer with large ventral process
(Fig. 55) fasciata (Osborn)

— Aedeagus without extra apodemal process;

pygofer with some type of apical process 2

2(1). Aedeagus with shaft laterally compressed, knife-
shaped, and with or without small lateral pro-
cesses (Figs. 10, 11, 14, 15) 3

'This paper is published with the approval of the Director of the Kentucky Agricultural Experiment Station as journal article No. 88-7-39.
Department of Entomology, University of Kentucky, Lexington, Kentucky 40546-0091.

Great Basin Naturalist Memoirs

No. 12

— Aedeagus with shaft tubular (Figs. 24, 2.5) . . 10
3(2). Style with a lateral subapical tooth (Fig. 4) . . . 4

— Style without subapical tooth 5

4(3). Pygofer truncate with a small apical process

(Fig. 1) decora (Fabricius)

— Pygofer gradually narrowing to bluntly
pointed apex, median of ventral margin with

two small teeth

tartessoides Linnavuori & Heller

5(3). Pygofer with apical process bifurcate (Figs.

13, 17, 18) boliviana (Osborn)

— Pygofer with apical process not bifurcate (Fig.
19) 6

6. Pygofer with apical process dorsally produced
and pointed (Fig. 5) 7

— Pygofer with apical process bird-head-shaped

(Fig. 9) 8

7. Style thickened to near apex (Fig. 8); aedea-
gus with small lateral processes (Figs. 6, 7) .
elongata. n. sp.

— Style narrowing to pointed apex (Fig. 46);
aedeagus with conspicuous processes (Figs.
44, 45) delicata, n. sp.

8. Aedeagus with conspicuous processes (Figs.
48, 49) ornata, n. sp.

— Aedeagus without processes (Figs. 10, 11) . . . 9

9. Style long and abruptly curved subapically

(Fig. 12) invita (Nast)

— Style short and evenly curved near apex (Fig.
22) lora, n. sp.

10(2). Style foot-shaped at apex (Fig. 26) 11

— Style pointed at apex (Fig. 42) 13

11(10). Aedeagus with shaft expanded at apex (Fig.

32) ahrupta, n. sp.

— Aedeagus with shaft not expanded at apex . . 12

12(11). Aedeagus with small apical processes (Fig. 24)

magna (Osborn)

— Aedeagus with larger subapical processes
(Fig. 28) expanda, n. sp.

13(10). Aedeagus with processes expanded at base

(Fig. 40) Columbiana (Nast)

— Aedeagus with thin, spinelike processes (Fig.
36) 14

14(13). Aedeagus with gonopore basal to processes

(Fig. 37) stigmata (Nast)

— Aedeagus with gonopore apical to processes
(Fig. 53) corvenda Kramer

Key to Known Females
1. Pronotum longitudinally banded 2

— Pronotum not so marked 3

2(1). Seventh sternum unusually long, over twice

length of sixth (Fig. 61) fasciata (Osborn)

— Seventh sternvmi shorter, about twice or less
length of sixth (Fig. 65) ynexicana, n. sp.

3(1). Seventh sternum truncate, with a nu'dian

emargination rngosa, n. sp.

— Seventh sternum not truncate, posterior mar-
gin usually rounded 4

4(3). Head with a frontal margin, bright yellow and

black marginata, n. sp.

— Head rounded to face 5

5(4). Ovipositor short, less than 2 mm in length

(Fig. 63) acuta, n. sp.

— Ovipositor long, more than 2 mm in length

(Fig. 62) ...6

6(5). Seventh sterninn with median emargination

of posterior margin V-shaped (Fig. 59)

ornata, n. sp.

— Seventh sternimi with median emargination
of posterior margin U-shaped (as in Fig. 62) .
decora (Fabricius)

The decora species group is characterized
by having orange transverse bands on the
head, and the aedeagus laterally flattened and
knife-shaped. The following seven species are
placed in this group.

Calliscarta decora (Fabricius)
Figs. 1-4, 66

Cicada decora Fabricius 1803: 69. (South America, possi-
bly from Brazil; Stockholm Museum).

Tettigonia decora: Signoret 1853: 327; Walker 1858: 200;
Dohrn 1859; 88.

Calliscarta decora: Stal 1869; 83; Dallas 1870: 495; Dis-
tant 1908; 524; Evans 1947; 230; Linnavuori 19.59:
29;Metcalfl967:946.

Thamnotettix magnificus Osborn 1924: 424. (Cuatro
Ojos, Santa Cruz, Bolivia; Carnegie Museum).
Ncti' synontjmtj

Idiotettix magnificus: Osborn 1929; 465; Oman 1938: 395;
Evans 1947: 192; Nast 19,52: 1; Maldonado
Capriles 19.54: 250; Metcalf 1966: 227.

Calliscarta magnifica: Linnavuori 1959: 28.

Idiotettix festivus Maldonado Capriles 19.54: 248.
(Cunucunuma River, F. T. Amazonas, Venezuela;
U.S. National Museum).

Length of males 9.5-11.5 mm.; female 12
mm. Head reddish yellow above antennal
sockets, yellow below, with three orange
transverse bands, one just above and one just
below ocelli and the other above base of an-
tennae. Pronotum reddish yellow with two
darker red transverse bands, one along ante-
rior margin and one across middle, and one
yellowish green band along posterior margin
continuing to anterior of lateral margins near
eyes. Scutellum reddish brown. Forewing
generally dark reddish brown, claval area
lighter brown with foiu- large, yellowish green
spots, costal margin red, and a row of si.\ to
eight large, yellowish green spots through
middle of wing from base to appendix, apex

1988

Freytag: Revision of Calliscarta

DECORA

ELONGATA

INVITA

Figs. 1-4. Calliscarta decora (Fabricius), from Peru; 1, lateral view of male genital segments; 2, lateral view of
aedeagus; 3, ventral view of aedeagus; 4, ventrolateral view of style.

Figs. 5-8. Calliscarta elongata, n. sp. , holotype; 5, lateral view of male genital segments; 6, lateral view of aedeagus;
7, ventral view of aedeagus; 8, ventrolateral view of style.

Figs. 9-12. Calliscarta invito (Nast), from Ecuador: 9, lateral view of male genital segments; 10, lateral view of
aedeagus; 11, ventral view of aedeagus; 12, ventrolateral view of style. All drawn to the same scale; the line equals 1
mm.

smoky brown. Ventral surface entirely yel-
low.

Male genitalia. — Pygofer truncate with a
short, apical, spinelike process (Fig. 1). Geni-
tal plates large, roundedly expanded at apex
(Fig. 1). Style long, bent at right angle three-
quarters of distance from base, apex pointed
with a subapical tooth on inner margin, and a
small but distinct spine at the bend (Fig. 4).
Aedeagus short, laterally flattened, broad in
lateral view, apex pointed, with a pair of small
processes on caudal margin near gonopore
(Figs. 2, 3).

Female genitalia. — Seventh sternum
large, twice length of sixth, with a shallow,
median, U-shaped emargination similar to or-
nata.

Type. — Holotype male of decora not seen,
but illustrated by Linnavuori (1959). Type is
from South America, and Dohrn (1859) gave
Brazil as the possible type locality.

Distribution. — Known are specimens
from: BOLIVIA: one female, Cuatro Ojos,
November 1913, Steinbach Coll. , (holotype of

magnificus) Carnegie Museum. Brazil: one
male, ? (holotype oi decora) Stockholm Mu-
seum. Ecuador: one male, Pompeya, 13-25
May 1965, Luis E. PeiiaColl., Ramos Collec-
tion; one male, Coca, May 1965, Luis E. Pena
Coll., Ramos Collection; two males, Napo,
Prov. Limoncocha, 10 June 1977, Dave L.
Vincent Coll., U.S. National Museum.
French Guyana: one male, Cabret Repub-
hque, 29 August 1975, Guyane Frangaise
Mission M. Boulard, et P. Pompanon Coll.,
Museum Paris. Guyana: one male, Bartica,
20-30 March 1901, H. S. Parish Coll., (allo-
type of magnificus) Ohio State University
Collection. Peru: four males, Avispas, Octo-
ber 1962, Luis E. Pena Coll., Ramos Collec-
tion. Venezuela: one male, Cunucunuma
River, F. T. Amazonas, 28 April 1950, J. Mal-
donado Capriles Coll. (holotype o( festivus)
U.S. National Museum; one male, Anacoco,
Bolivar, 60 M, 6°5'N:6r8'W, 10-30 August
1979, Expedicion La Salle, University of Cen-
tral Venezuela Collection.

Notes. — This species appears to be widely

Great Basin Naturalist Memoirs

No. 12

distributed in South America but is poorly
represented in collections. Much confusion
concerning the identification of this species
has occurred because the color pattern is not
fully developed in some specimens. How-
ever, this species can be quickly recognized
by the small spine on the male style, which is
found on only one other known species,
tartessoides. For now the female must be rec-
ognized by the color pattern, which is the
same as in the male.

The types of magnificus and festivus were
examined and found to represent the same
species. Linnavuori (1959) synonymized fes-
tivus with decora.

Calliscarta tartessoides Linnavuori & Heller

Calliscarta tartessoides Linnavuori & Heller 1961: 3.
(Hacienda Mascoitania, Peru; Staatlichen Mu-
seum fur Naturkunde, Stuttgart).

This species was illustrated when de-
scribed, and no additional information can be
added at this time. It appears to be closely
related to decora.

Length of male 9.5 mm; female unknown.
Brownish yellow. Head light yellow, with a
reddish orange transverse band across vertex
between eyes; face with an orange transverse
band between ocelli and another across mid-
dle of face, and a dark brown spot underneath
each ocellus. Pronotum with anterior margin
light yellow, posterior two-thirds darker mar-
bled with darker orange. Scutellum faintly
dark orange. Forewing mostly translucent,
with veins brown; costal vein darker brown in
middle. Ventral surface light yellow with
some yellow-orange and brown markings on
legs and abdomen.

Male genitalia. — Pygofer narrowing to a
posteriorly projecting, bluntly pointed apex,
with two small spines on ventral margin near
middle. Genital plates large and truncate at
apex. Style long, bent laterally on apical
fourth, apex pointed, a small subapical tooth
on inner margin. Aedeagus laterally flattened,
knife-shaped, without noticeable processes,
gonopore on ventral margin near middle of
shaft.

Type. — Holotype (male), Peru, Hacienda
Mascoitania, 10-12 November 1957, Urwald
Coll., Staatlichen Museum fiir Naturkunde,
Stuttgart.

Notes. — This species is known only from
the type specimen, which I have not seen. It is

a species like decora, with a tooth on the style,
but a color pattern that is more like magna.
More material is needed to properly place this
species.

Calliscarta elongata, n. sp.

Figs. 5-8

This species resembles decora in general
appearance, but with different male genitalia.

Length of males 10-10.1 mm; female un-
known. Head creamy yellow, crown having a
red-violet tinge, with four orange transverse
bands, one across middle of crown between
eyes, one just below ocelli, two fainter bands
across postclypeus; a black band on each side
from base of antenna to anteclypeus along side
of postclypeus. Pronotum orange, with a band
along posterior margin and two central spots
greenish yellow. Scutellum orange with a
small yellow spot in middle of each lateral
margin. Forewing orange anteriorly, fading to
brown posteriorly, median of costal vein with
a reddish tinge, four greenish yellow spots on
clavus, three at base and one near middle,
four yellow spots on corium, apical cells
smoky brown with an L-shaped translucent
spot pattern. Ventral surface generally
creamy yellow, fore tibiae and tarsi brownish
yellow.

Male genitalia. — Pygofer narrowing to
apex, apex a dorsally projecting, pointed pro-
cess (Fig. 5). Genital plate enlarged and ex-
panded on apical half, apex bluntly rounded
(Fig. 5). Style stout, uniform width through-
out, hooked apically, apex with a very small
subapical tooth (Fig. 8). Aedeagus laterally
flattened, knife-shaped, gonopore medially
on ventral margin, a pair of small lateral pro-
cesses basal to gonopore (Figs. 6, 7).

Type. — Holotvpe (male), Ecuador, Napo,
Lago Agrio (18 km E), 28 August 1975, Rio
Aguarico, collected on gravel bar at black-
light, Langley, Cohen, Cantor, Yingling
Coll. , Ecuador-Peace Corps-Smithsonian In-
stitution Aquatic Insect Survey, in the U.S.
National Museum. Paratypes: Ecuador, one
male, same data as holotype, except 23 Sep-
tember 1975 and Andrea Langley Coll. , Uni-
versity of Kentucky Collection; one male,
same data as holotype, except 19 September
1975 and Andrea Langley Coll., U.S. National
Museum; Peru, Avispas, October 1962, Luis
E. PenaColl., Ramos Collection.

1988

Freytac: Revision of Calliscarta

Calliscarta invita (Nast)
Figs. 9-12

Idiotettix invitus Nast 1952: 3. (Santa Inez, Ecnador,
Polish Museimi of Zoology); Maklonado Capriles
1954: 250; Metcalf 1966: 226.

Calliscarta invita: Linnavuori 1959: 30; Linnavuori and
Heller 1961: 3.

Length of males 10-10.6 mm; female un-
known. Head brownish yellow, with a tinge of
violet on crown and postclypeus; an orange
transverse band across middle of crown be-
tween eyes, another orange transverse band
below ocelli, two brownish orange transverse
bands on postclypeus, a black band on each
side from base of antennae to anteclypeus
along sides of postclypeus. Pronotum mostly
orange brown, lateral margins yellow. Scutel-
lum light brown. Forewing light brown, with
veins and obscure spots darker brown, me-
dian of costa reddish orange, small, white,
translucent spots on clavus and corium; apical
area smoky brown with three translucent
spots. Ventral surface mostly brownish yel-
low, brown spot on propleura and sterna, fore
legs with most of tibiae and entire tarsi brown,
middle legs with base and apex of tibiae and
entire tarsi brown.

Male genitalia. — Pygofer truncately nar-
rowed to apex, which is projected slightly dor-
sad in the shape of a bird's head (Fig. 9).
Genital plate large, expanded in lateral view,
truncate (Fig. 9). Style elongate, even width
throughout, hooked apically, with a pointed
apex (Fig. 12). Aedeagus laterally flattened,
knife-shaped, without processes, gonopore on
ventral margin near base of shaft (Figs. 10,
11).

Type. — Holotype (male), Ecuador, Santa
Inez, R. Haensch Coll., Polish Museum of
Zoology.

DiSTRlRUTlON. — Five males are known for
this species. Linnavuori and Heller (1961) re-
ported two males from Peru, Hacienda Mas-
coitania, 10-12 September 1957, Urwald
Coll., Staatlichen Museum fiir Naturkunde,
Stuttgart. I have seen two males from Ecua-
dor, Rio Margajita, Rio Pastaza, 1250 m, 20
March 1939, F. M. and H. H. Brown Coll.,
University of Kansas Collection.

Calliscarta boliviana (Osborn)
Figs. 13-18

Idiotettix bolivianus Osborn 1929: 466. (Coroico, La Paz,
Bolivia, Ohio State University Collection); Oman

1938: 395; Nast 1952: 1; Maldonado Capriles 1954:

250; Metcalfl966: 225.
Calliscarta boliviana. Linnavuori 1959: 29.
Calliscarta ting,a Kramer 1963: 205. (Tingo Maria,

Iluanuco, Peru, U.S. National Museum). NeuJ

sifnoniimij

Length of males 9.1-10 mm; female un-
known. General color as in invita, pronotum
with a band of yellow along posterior margin
and lateral margins and white, translucent
spots on forewings less conspicuous.

Male genitalia. — Pygofer quite variable,
narrowing to near apex, which projects dorsad
in some type of bifurcate process (Figs. 13, 17,
18). Genital plates greatly expanded on apical
half (Fig. 13). Style elongate, narrowing
slightly to apex, hooked apically, apex with a
ventral subapical tooth (Fig. 16). Aedeagus
laterally flattened, knife-shaped, gonopore on
ventral margin near base of shaft, with a small
pair of laterally projecting processes basal to
gonopore (Figs. 14, 15).

Type. — Holotype (male), Bolivia, Coroico,
Ohio State University Collection.

Distribution. — This species is known from
10 male specimens. Besides the two types, I
have seen: BOLIVIA, one male, Cristal Mayn,
Chapare, 28 August 1949, L. E. Peiia Coll.,
University of Kansas Collection; Peru, four
males, Avispas, September 1962, Luis E.
Pefia Coll.; two males, Quincemil, August
1962, Luis E. Peria Coll., Ramos Collection,
and one from each locality in University of
Kentucky Collection; Panama, El Volcan de
Chiriqui, 26 February 1936, W. J. Gertsch
Coll., American Museum of Natural History.

Notes. — This species is quite variable in
the shape of the apex of the pygofer, but it is
the only species of the genus with the apex
bifurcate. No two specimens seen have the
same shape, and I consider this to be just a
variation at this time; however, this may
prove to be a complex of very closely related
species.

Calliscarta lora, n. sp.

Figs. 19-22

This species resembles invita in general ap-
pearance, but with distinct male genitalia.

Length of male 9.3-9.9 mm; female un-
known. Head, pronotum, and scutellum as in
elongata. Forewing with clavus mostly or-
ange, with three yellow spots, two transverse
spots anteriorly, one longitudinal spot along
posterior part of claval suture; corium brown.

Great Basin Naturalist Memoirs

No. 12

14 15

BOLIVIANA

Figs. 13-16. Calliscarta boliviana (Osborn), liolotype: 13, lateral view of male genital segments; 14, lateral view of
aedeagus; 15, ventral view of aedeagus; 16, ventrolateral view of style.

Figs. 17—18. Lateral views of male genital segments of other specimens of C. boliviana from Bolivia.

Figs. 19-22. Calliscarta lora, n. sp., holotype; 19, lateral view of male genital segments; 20, lateral view of aedeagus;
21, ventral view of aedeagus; 22, ventrolateral view of style. All drawn to the same scale; the line equals 1 mm.

median of costal vein with a reddish tinge,
with four yellow spots, two at base, one (some-
times two) above reddish tinge of costa, and
one a continuation of longitudinal spot on
clavus; apical cells smoky brown, with three
small, translucent spots. Ventral surface
creamy yellow, fore tibiae and tarsi brownish
yellow.

Male genitalia. — Pygofer narrowing to
apex that extends as a large bird's-head-
shaped process (Fig. 19). Genital plate gradu-
ally expanding to a large, truncate apex (Fig.
19). Style elongate, hooked apically, with a
pointed apex (Fig. 22). Aedeagus laterally flat-
tened, knife-shaped, gonopore on ventral
margin near middle, without processes (Figs.
20, 21).

Type. — Holotype (male), Peru, Quincemil,
August 1962, Luis E. Pena Coll., in the
Ramos Collection but will be deposited in
U.S. National Museum through courtesy of
Dr. Ramos. Paratypes: Peru, two males,
same data as holotype; one male, Quincemil,
October 1962, Luis E. Peiia Coll.; two males,
Avispas, September 1962, LuisE. Pena Coll.,
three in the Ramos Collection and two in the
University of Kentucky Collection.

Notes. — This species has a distinct color
pattern and male genitalia that make it possi-

ble to separate it from other known species
that also occur in Peru.

Calliscarta delicata.
Figs. 43-46

n. sp.

This species resembles invita in general ap-
pearance, but with distinct male genitalia.

Length of male 8.5-10.1 mm; female
unknown. Head yellow, with four orange
transverse bands, one above ocelli, one be-
low ocelli, one between antennae and one
along lower margin of postclypeus. Pronotum
orange, with green posterior margin extend-
ing along lateral margins and a green trans-
verse band in anterior half Scutellum mostly
orange. Forewing generally smoky brown,
with claval suture, anterior third of costa, and
small spots at ends of claval veins green; costa
red near middle; corium with two orange
bands extending from near base to apical
veins, one along claval suture, other along
costa; apical cells smoky brown with three
transverse, opaque spots. Abdominal seg-
ments orange.

Male genitalia. — Pygofer narrowing to
somewhat expanded apex, with a dorsally pro-
jecting, apical, pointed process (Fig. 43).
Genital plate gradually expanded to a large,
truncate apex (Fig. 43). Style narrowing and

1988

FreytaG: Revision of Calliscarta

MAGNA

28 29

EXPANDA

Figs. 23-26. Calliscarta nuigna (0.shorn), from Brazil: 23, lateral view of male genital segments; 24, lateral view of
aedeagus; 25, ventral view of aedeagus; 26, ventrolateral view of style.

Figs. 27-30. Calliscarta expanda. n. sp., holotype: 27, lateral view of male genital segments; 28, lateral view of
aedeagus; 29, ventral view of aedeagus; 30, ventrolateral view of style.

Figs. 31-34. Calliscarta abrupta, n. sp., holotype; 31, lateral view of male genital segments; 32, lateral view of
aedeagus; 33, ventral view of aedeagus; 34, ventrolateral view of style. All drawn to the same scale; the line equals 1

curving laterally near pointed apex (Fig. 46).
Aedeagus laterally compressed with a pair of
long, curved processes from base of shaft,
gonopore medially on ventral margin, apex
bluntly pointed (Figs. 44, 45).

Type. — Holotype (male), Peru, Tingo
Maria, 20 June 1982, Fisk, Ohio State Univer-
sity Collection. Paratypes: two males, same
data as holotype, one in Ohio State University
Collection and one in University of Kentucky
Collection. Two males, Bolivia, El Palmar,
Cochabamba, Chapare, 8-15 September
1955, Luis E. PeiiaColl., Ramos Collection.

Notes. — This species can be separated
from invita by the longer processes on the
aedeagus.

The ornata species group is characterized
by a single, transverse, black band on the face
that replaces the orange transverse bands of
the decora group, and by an aedeagus that is
flattened and knife-shaped. Only one species,
ornata, is placed here.

Calliscarta ornata, n. sp.
Figs. 47-50, 59, 67

This species resembles invita in general ap-
pearance, but with unique black band on face
instead of usual orange bands.

Length of male 9.4-10.1 mm; female 10-
11 mm. Head creamy yellow with a reddish
tinge; an irregular, black, transverse band
below ocelli, a large, black spot in middle of
postclypeus, a black, V-shaped band from an-
tennal bases along sides of postclypeus over
entire anteclypeus. Pronotum creamy yellow,
with a large, irregular, transverse, brown
band across middle, posterior margined with
greenish yellow. Scutellum mostly creamy
yellow. Forewing brown, with many large,
greenish yellow spots, apical cells smoky
brown, with two large, translucent spots.

Male genitalia. — Pygofer narrowed to a
truncate apex, with a large, nearly bird's-
head-shaped apical process (Fig. 47). Genital
plate large, greatly expanded at apex, which is
bluntly rounded (Fig. 47). Style elongate,

Great Basin Naturalist Memoirs

No. 12

STIGMATA

COLUMBIANA

44 45

DELICATA

Figs. 35-38. Calliscarta stigmata (Nast), from Colombia: 3.5, lateral view of male genital segments; 36, lateral view of
aedeagus; 37, ventral view of aedeagus; 38, ventrolateral view of style.

Figs. 39-42. Calliscarta columbiana (Nast), from Colombia; 39, lateral view of male genital segments; 40, lateral
view of aedeagus; 41, ventral view of aedeagus; 42. ventrolateral view of style.

Figs. 43-46. Calliscarta delicata, n. sp., holotype; 43, lateral view of male genital segments; 44, lateral view of
aedeagus; 45, ventral view of aedeagus; 46, ventrolateral view of style. All drawn to the same scale; the line equals 1

hooked apically, with a pointed apex (Fig. 50).
Aedeagus laterally flattened, knife-shaped,
with a pair of recurved processes on ventral
margin near base of shaft, gonopore on ventral
margin just apical to bases of processes (Figs.
48, 49).

Female genitalia. — Seventh sternum en-
larged, rounded posterior margin with a
large, median, V-shaped emargination (Fig.
59). Ovipositor extending twice its width be-
yond pygofer. Pygofer narrowing to apex in
ventral view.

Type. — Holotype (male), French Guyana,
Saint-Elie, 30 August-2 September 1975,
Guyane Franyaise Mission, M. Boulard et P.
Pompanon Coll., Paris Museum. Allotype
(female), same data as holotype. Paratypes:
French Guyana, one female, Garbet Lavaud
(Rive Surinamienne), 24-26 September 1975,
Itani (Guyanes) Mission, M. Boulard, P. Jauf-
fret et P. Pompanon Coll.; one female, Em
Gouchure de la Crique Oyaricoulet, 23 Sep-
tember 1975, Itani (Guyanes) Mission, M.
Boulard, P. Jauffret et P. Pompanon Coll.;
one female, Alicoto-Oyapock-Guyane, 13
November 1969, Piege lumineux, Guyane
Mission, Balachowsky, Gruner Coll., Oct.-

Nov. 1969, Paris Museum; Brazil, one fe-
male, Chapada, September, C. F. Baker
Coll.; one female, Chapada, October, C. F.
Baker Coll., U.S. National Museum; Vene-
zuela, one male, San Carlos de Rio Negro,
T. F. Amazonas, 7-13 September 1982, A.
Chacon, G. Yepez Gil Coll.; one female, Sta.
Lucia, T. F. Amazonas, 15-21 September
1982, A. Chacon, G. Yepez Gil Coll.; one
male, one female, Rio Surukum, Carretera
Sta., Elena Icabaru, Bolivar, 850 m, 19-31
January 1985, F. Fernandez Y., Anibal
Chacon, Jurg Demarmels Coll., University of
Central Venezuela Collection.

Notes. — This species can be easily sepa-
rated from the known species of this genus on
the basis of the dark band across the face.

The magna species group is characterized
by having the head mottled with brown and
the aedeagal shaft tubular with apical or sub-
apical processes. The following three species
are placed in this group.

Calliscarta magna, (Osborn)

Figs. 23-26

Idiocerus niap,nus Osborn 1923: 13. (San Antonio de Gua-
pore, MattoGrosso, Brazil, Carnegie Museum).

1988

Freytag: Revision of Calliscarta

48 49

OR N ATA

52 53

50 CORVENDA

Figs. 47-50. Calliscarta oniata, n. sp., holotype: 47, lateral view of male genital segments; 48, lateral view of
aedeagus; 49, ventral view of aedeagus; 50, ventrolateral view of style.

Figs. 51-54. Calliscarta corvenda Kramer, from Venezuela: 51, lateral view of male genital segments; 52, lateral
view of aedeagus; 53, ventral view of aedeagus; 54, ventrolateral view of style.

Figs. 55-58. Calliscarta fasciata (Osborn), from Brazil: 55, lateral view of male genital segments; 56, lateral view of
aedeagus with connector to anal tube; 57, ventral view of aedeagus and connector to anal tube; 58, ventrolateral view of
style. All drawn to the same scale; the line equals 1 mm.

Idiotettix magnus: Maldonado Capriles 1954: 247; Met-

calf 1966: 227.

Calliscarta rrmgna: Linnavuori 1959: 30.

Length of male 9-10 mm; female unknown.
Head, pronotum, and scutellum creamy yel-
low with a reddish violet tinge, greatly mot-
tled with brown. Forewing mostly translu-
cent, veins dark brown; clavus with a large,
fuscous spot at middle and apex; apical cells
smoky brown, with two transverse, translu-
cent spots. Ventral surface mostly creamy yel-
low, with fore and middle legs with apex of
tibiae and entire tarsi brownish yellow.

Male genitalia. — Pygofer gradually nar-
rowing to a pointed apex (Fig. 23). Genital
plate large, greatly expanded on apical half
(Fig. 23). Style elongate, hooked apically,
apex bluntly pointed (Fig. 26). Aedeagus
tubular, with pair of small, lateral, apical pro-
cesses, gonopore subapical on ventral margin
(Figs. 24, 25).

Type. — Holotype (male), Brazil, Rondo-
nia, Santo Antonio de Guapore (Rio Gua-

pore), 26 July 1909, Haseman Coll., Carnegie
Museum.

Distribution. — Known only from Brazil on
the basis of the type and the following speci-
mens: Brazil, five males, Sao Felice, Camp
IV, Moyen Xingu, (Bresil) Mission, M. Bou-
lard, P. Jauffret et P. Pompanon Coll. , four in
Museum Paris and one in University of Ken-
tucky Collection.

Calliscarta expanda, n. sp.

Figs. 27-30

This species resembles magna in general
appearance, but with distinct male genitalia.

Length of male 9.7-10.5 mm.; female un-
known. Head, pronotum, and scutellum
creamy violet, mottled with brown. Forewing
brown, with most cells translucent, veins dark
brown; apical cells smoky brown, with three
or four large, translucent, transverse spots.
Ventral surface creamy yellow heavily marked
with brown.

Male genitalia. — Pygofer narrowing to a

Great Basin Naturalist Memoirs

No. 12

pointed apex (Fig. 27). Genital plate large,
expanded on apical half (Fig. 27). Style elon-
gate, hooked apically, with apex bluntly
pointed, notched subapically (Fig. 30). Aede-
agus tubular, with subapical, large, lateral
pair of processes, gonopore on ventral margin
subapical to processes (Figs. 28, 29).

Type. — Holotype (male), Ecuador, Napo,
Lago Agrio, 19 September 1975, at black-
light, Andrea Langley Coll., Ecuador-Peace
Corps-Smithsonian Institution Aquatic In-
sect Survey, U.S. National Museum. Para-
types: Ecuador, one male, same data as holo-
type, except 23 September 1975, (18 km E)
Rio Aguarico; two males, same data, except 28
August 1975, collected on gravel bar, Lang-
ley, Cohen, Cantor, Yingling Coll., U.S. Na-
tional Museum; Peru, one male, Loreto,
Ucayali R. , Yarina Cocha, 24 September
1953, Peter Hocking Coll., Field Museum;
one male, Quincemil, August 1962, Luis E.
Peria Coll. ; one male, Loromavo, September
1962, Luis E. Peiia Coll., Ramos Collection;
Bolivia, Rurrenbaque, 10-23 October 1956,
Luis E. Peiia Coll., Ramos Collection.

Notes. — This species is similar to magna,
but the aedeagus is larger and with longer,
more subapical processes.

Calliscarta abnipta, n. sp.

Figs. 31-34

This species is related to magna, but with a
distinctly different aedeagus.

Length of male 9.8 mm; female unknown.
Head, pronotum, and scutellum mostly yel-
low tinted with red. Head with a small, brown
spot behind each ocellus next to eye, another
larger, brown spot below each ocellus next to
eye. Postclypeus with brown dashes on each
side and brown speckles over remainder of
surface. Pronotum speckled with brown.
Forewings mostly brown, claval area with
opaque spots at ends of claval veins, costal
vein yellow, two cell rows behind costal vein
transparent, giving wing appearance of having
a brown longitudinal band through middle,
apical cells smoky brown with several large,
opaque spots.

Male genitalia. — Pygofer narrowing grad-
ually to an upturned, pointed apex (Fig. 31).
Genital plate enlarged on apical half, truncate
(Fig. 31). Style nearly same width to apex,
hooked apically, with apex foot-shaped (Fig.
34). Aedeagus tubular, with shaft bent at right

angle near middle, apex expanded with a pair
of subapical large processes extending along
sides of shaft halfway to base, gonopore on
ventral margin just basal to processes (Figs.
32, 33).

Type, — Holotype (male), Bolivia, Rurren-
abaque, 10-23 October 1956, Luis E. Peiia
Coll., in Ramos Collection but is being de-
posited in U.S. National Museum through
courtesy of Dr. Ramos.

Notes. — This species can be separated
from the other species of the genus by the
distinct aedeagus.

The stigmata species group is characterized
by having the head mottled with brown, and
the aedeagal shaft tubular with processes near
middle. The following four species are placed
in this group.

Calliscarta stigmata (Nast)
Figs. ,3.5-38

Idiotettix stigmatus Nast 1952: 2. (Hac. Pehlke [probably
Magdalena], Colombia, Polish Museum of Zool-
ogy); Maldonado Capriles 19.54: 250; Metcalfl966:
227.

Calliscarta stigmata: Linnavuori 1959: 31.

Length of male 8.5-9.4 mm; female un-
known. Head, pronotum, and scutellum gray-
ish yellow tinged with violet, mottled with
brown; orange bands of crown and face ob-
scure. Forewing brown, with numerous
whitish, translucent spots. Ventral surface
brownish yellow; legs banded with brown.

Male genitalia. — Pygofer narrowing to a
long, tubular, apical area, apex with small,
dorsally projecting process (Fig 35). Genital
plate long, expended on apical half, bluntly
rounded at apex (Fig. 35). Style long, bent
apically, with pointed apex (Fig. 38). Aedea-
gus tubular, shaft elongate, apex bent dorsad
and pointed, a pair of lateral processes at base
of apical bend, gonopore basal to processes on
ventral margin (Figs. 36, 37).

Type. — Holotvpe (male), Colombia, Ha-
cienda Pehlke, ' 1925, Pehlke Coll., Polish
Museum of Zoology.

Distribution. — This species is known from
eight male specimens. The holotype and five
paratypes are from Colombia and are in the
Polish Museum of Zoology, and I have seen
one male, Colombia, Alban, Cund., 10 Sep-
tember 1965, J. A. Ramos Coll., Ramos Col-
lection; and one male, Venezuela, Rio Frio,

1988

Freytag: Revision of Calliscarta

ORNATA

MARGINATA

FA SCI ATA

Figs. 59-61. Calliscarta spp., ventral view offemale segments: 59, C. ornata, n. sp., allotype; 60, C. nuirg,inata, n.
sp., holotype; 61, C.fasciata (Osborn), from Brazil. All drauni to the same scale; the line equals 1 mm.

Tachira, 600 m, 11-14 December 1980, J. A.
Clavijo, A. Chacn, J. AyalaColl., University
of Central Venezuela Collection.

Calliscarta columbiana (Nast)
Figs. 39-42

Idiotettix columbianns Nast 1952: 2. (Hac. Pehlke [proba-
bly Magdalena], Colombia, Polish Museum of
Zoology); Maldonado Capriles 1954: 250; Metcalf
1966: 226.

Calliscarta columbiana: Linnavuori 1959: 29.

Length of male 8-9.1 mm; female un-
known. General color pattern as in stigmata,
except lighter in color and legs only faintly
banded with brown.

Male genitalia. — Pygofer narrowing to
truncate apex with a small, dorsad projecting
process (Fig. 39). Genital plates large, ex-
panded, truncate at apex (Fig. 39). Style elon-
gate, hooked apically, with pointed apex
(Fig. 42). Aedeagus semitubular, apex bluntly
pointed, with a pair of basally expanded, lat-
eral processes halfway to base, gonopore on
ventral margin at base of processes (Figs. 40,
41).

Type, — Holotype (male), Colombia, Ha-
cienda Pehlke, 1921, Pehlke Coll., Pohsh
Museum of Zoology.

Distribution. — This species is known only

from Colombia from eight male specimens.
The holotype and six paratypes are in the
Polish Museum of Zoology, and I have seen
one male, Alto Rio Opon, Santander, January
1950, L. RichterColl., Ramos Collection.

Calliscarta corvenda Kramer

Figs. 51-54

Calliscarta corvenda Kramer 1963: 210. (Covendo, La
Paz, Bolivia, U.S. National Museum).

Length of male 9-9.8 mm.; female un-
known. Color pattern similar to stagmata, ex-
cept spots of forewings not forming rows or
bands.

Male genitalia. — Pygofer narrowing to
inwardly hooked and pointed apex (Fig. 51).
Genital plate large, expanded on apical half
(Fig. 51). Style very elongate, hooked api-
cally, with pointed, upturned apex (Fig. 54).
Aedeagus semitubular, pair of lateral pro-
cesses on ventral margin near middle of shaft,
gonopore subapical on ventral margin (Figs.
52, 53).

Type. — Holotype (male), Bolivia, Coven-
do, August 1921, Mulford Biol. Expedition,
Wm. M. Mann Coll., U.S. National Museum.

Distribution. — Four specimens are
known, including the type and a paratype from

Great Basin Naturalist Memoirs

No. 12

RUGOSA

ACUTA

MEXICANA

Figs. 62-65. Calliscarta spp. , ventral view of female segments: 62, C. rugosa, n. sp. , holotype; 6.3, C. acuta, n. sp. ,
holotype; 64, C acuta, n. sp., paratype; 65, C. mexicana, n. sp., holotype. All drawn to the same scale; the line equals
1 mm.

Bolivia. Other specimens: Venezuela, two
males, Barinitas, 15 km SW Barinas, 25 Feb-
ruary 1969, Duckworth and Dietz Coll., one
in U.S. National Museum and one in Univer-
sity of Kentucky Collection.

Calliscarta rugosa, n. sp.

Figs. 62, 68

Resembling columbiana in general color
pattern, but with lighter, more reddish brown
markings.

Length of female 10. 1 mm; male unknown.
General color pattern as in columbiana, but
with forewing darker and more reddish
brown.

Female genitalia. — Seventh sternum large,
truncate, with a large, median, U-shaped
emargination (Fig. 62). Ovipositor extending
its length beyond pygofer.

Type. — Holotype (female), Venezuela,
Aragua, El Limon, 450 m, 23 August 1979,
LuzdeMercurio, F. Fernandez Y. Coll., Uni-
versity of Central Venezuela Collection.

Notes. — This female may be the female of
columbiana or corvenda. However, since it
was collected along the northern mountains of
Venezuela, it is just as likely to represent a
separate species. The color pattern is suffi-
ciently different from the known species of the
stigmata species group to warrant its descrip-
tion at present.

The following two species are not placed in
a species group at this time, as the males are
unknown and the color patterns give little
indication as to how these relate to other
known species. However, they are closer to
the preceding groups than to the group fol-
lowing them.

Calliscarta marginata, n. sp.
Figs. 60, 69

This species resembles ornata, but with a
brighter color pattern and the anterior margin
of the crown more distinct.

Length of female 10.5 mm; male unknown.
Head creamy yellow with a U-shaped band in
middle of crown, a wider U-shaped band be-
hind eyes over ocelli and along anterior mar-
gin, two transverse bands between antennae
and ocelli, two dashes in middle of lower part
of postclypeus, and a V-shaped band from
underneath eyes along lateral margins of post-
clypeus down middle of anteclypeus, black.
Pronotum black, with a yellow longitudinal
band down median; also a large, yellow spot
on each side with a smaller cream spot near
anterior margin between yellow spot and
band. Scutellum black, with cream spots.
Forewing black, with six large, yellow spots or
dashes on clavus, eight large, yellow spots or
dashes on corium, and four smaller cream
spots in apical cells.

1988

FreytaG: Revision of Calliscarta

(

..Hill. TIUp*^

Figs. 66-71. Calliscarta spp., lateral aspect: 66, C. decora (Fabricius), male from Peru; 67, C. ornata, n. sp., male
paratype from French Guyana; 68, C. rugosa, n. sp. , female holotype; 69, C. marginata, n. sp. , female holotype; 70, C.
acuta, n. sp., female holotype; 71, C mexicana, n. sp., female holotype.

Female genitalia. — Seventh sternum
large, with posterior margin with a median,
V-shaped emargination (Fig. 60). Ovipositor
extending beyond pygofer three times its
width.

Type. — Holotype (female), Honduras,
Com., Lago Yojoa, 19 July 1974, C. W. and
L. B. O'Brien and Marshall Coll., the Ohio
State University Collection.

Notes. — This species is quite distinct from
other members of the genus both in color
pattern and in having a slight anterior margin
of the head.

Calliscarta acuta, n. sp.

Figs. 63, 64, 70

Resembling marginata, this species is

smaller and has a distinctly different color
pattern.

Length of female 8.9-9 mm; male un-
known. Head creamy yellow, with a spot on
either side of median near posterior margin of
crown, a spot on either side of median on
anterior margin of crown, an area around an-
tennae down lateral margins of postclypeus,
lora, and anteclypeus black. Pronotum
creamy yellow with a large spot either side of
median. Scutellum black with a median, or-
ange spot. Forewing green, with three black
bands, one at end of scutellum, one across
middle, and one at end of clavus, apical cells
black.

Female genitalia. — Seventh sternum

Great Basin Naturalist Memoirs

No. 12

large, roundedly produced, with a small, me-
dian, V-shaped emargination (Fig. 63).
Ovipositor extending beyond pygofer nearly
twice its width.

Type. — Holotype (female), Venezuela,
T. F. Amazonas, 20 December 1981, eacuri,
G. Yepez Gil Coll., University of Central
Venezuela Collection. Paratype (female),
Peru, Quincemil, August 1962, Luis E. Peria
Coll., Ramos Collection.

Notes. — The striking color pattern makes
this species easy to identify. The two speci-
mens differ slightly in color pattern and in the
shape of the female genitalia, but this is proba-
bly individual variation as they come from
quite different localities.

The fasciata species group is characterized
by the longitudinal bands on the pronotum
and the unusual, extra apodemal process on
the aedeagus. The following two species be-
long to this group.

Calliscarta fasciata (Osborn)
Figs. 55-58, 61

Idiotettix fasciatus Osborn 1929: 466. (Hohenau, Itapua,
Paraguay, Ohio State University Collection);
Oman 1938: 395; Evans 1947: 190;' Nast 1952: 1;
Maldonado Capriles 19.54: 248; Metcalf 1966: 226.

Calliscarta fasciata: Linnavuori 1959: 30.

Idiotettix brunnetis Osborn 1929: 467. (Hohenau, Itapua,
Paraguay, Ohio State University Collection);
Oman 1938: .395; Nast 1952: 1; Maldonado
Capriles 1954: 248; Metcalf 1966: 226. Neiv syn-
onijmij

Calliscarta hrunnca: Linnavuori 1959: 32.

Idiotettix laiitus Nast 1952: 3. (Porto Epitacio, Sao Paulo,
Brazil, Polish Museum of Zoology); Maldonado
Capriles 19.54: 248; Metcalf 1966: 227.

Length of male 8-9.5 mm; female 9.8 mm.
Head yellow, with a transverse band above
ocelli, one below ocelli, a spot on each anten-
nal ledge, a pair of longitudinal bands on post-
clypeus, and much of gena and lora orange;
a row of brown or black transverse lines on
each side of postclypeus. Pronotum yellow,
with anterior margin, four wide, longitudinal
bands, and lateral margins orange. Scutellum
yellow, with orange-brown longitudinal band
down median. Forewings orange fading to
smoky brown at apex; claval and discal veins
bordered with translucent puri:)le bordered
with dark brown, apical half with large, irreg-
ularly shaped white spots bordered with
brown, apical cells with whitish, translucent
spots. Ventral surface mostly yellow.

Male genitalia. — Pygofer large, tnmcate.

with a long process from ventral margin (Fig.
55). Genital plate large, expanded apical half,
bluntly rounded at apex (Fig. 55). Style short,
slightly curved laterad, with a pointed apex
(Fig. 58). Aedeagus stout, curved dorsad api-
cally, with a pair of lateral flanges from base of
shaft to near apex, extra apodeme at base con-
necting aedeagus to anal tube (Figs. 56, 57).

Female genitalia. — Seventh sternum
greatly enlarged, rounded on posterior mar-
gin with a V-shaped, median emargination
(Fig. 61). Ovipositor extending beyond
pygofer its own width. Pygofer narrowed on
apical third.

Type. — Holotype (male), Paraguay, Ho-
henau, Ohio State University Collection.

Distribution. — This species is known from
several hundred specimens from Brazil,
Paraguay, and Peru. The only known female is
in the University of Kansas Collection.

Notes. — Linnavuori (1959) placed lautus as
a synonym of fasciatus. The type of brunnea
represents the teneral form of this species;
consequently, it is placed in synonymy.

Calliscarta mexicana, n. sp.

Figs. 65, 71

This species resembles fasciata, but it is
smaller and the apical area of the forewing is
without large, transparent spots.

Length of female 8 mm; male unknown.
Head creamy yellow, with a large, orange
transverse band across middle of crown, an
orange transverse band just below ocelli with
ends below ocelli dark brown, an orange-
brown transverse band across postclypeus
halfway between antennae and ocelli with ex-
tensions down both sides of postclypeus as a
brown longitudinal band to anteclypeus, and a
dark brown area around bases of antennae
extending along lateral margins of post-
clypeus. Pronotum creamy yellow with four
wide, orange-brown longitudinal bands, me-
dian pair extending as brown longitudinal
bands on scutelhuu. Forewing orange-brown
with longitudinal bands of creamy white, one
along costa, one along claval vein on clavus,
one along commissure, as well as white areas
along apical longitudinal veins.

Female genitalia. — Seventh sternum
slightly enlarged, with posterior margin
roundly produced with a slightly rounded me-
dian emargination (Fig. 65). Ovipositor ex-
tending only slightly more than its width be-
yond pygofer.

1988

Freytag: Revision of Calliscarta

Type. — Holotype (female), Mexico, Tama-
zunchale, S. L. Potosi, 19 June 1941, Henry
S. Dybus Coll., Field Museum.

Notes. — This species is very similar to fas-
ciata in overall color pattern. These two spe-
cies form a species group that is quite different
from the other species of this genus.

Acknowledgments

Material for this study has been gathered
over the last 20 years from the institutions
listed below. I certainly thank the curators for
their patience over these long years, for the
loan of these most valuable specimens, and
also for the opportunity to study type mate-
rial: American Museum of Natural History,
New York City, Dr. Jerome G. Rozen, Jr.,
and Dr. Randall T. Schuh; Carnegie Mu-
seum, Pittsburg, Dr. Chen W. Young; Field
Museum of Natural History, Chicago, Dr. R.
Wenzel; Ohio State University, Columbus,
Dr. Charles A. Triplehorn; Museum National
d'Histoire Naturalle, Paris, Dr. Michel Bou-
lard; Ramos Collection, Mayaguez, Dr. J. A.
Ramos; United States National Museum,
Washington, D.C., Dr. James P. Kramer;
University of Kansas, Lawrence, Dr. George

W. Byersand Dr. Peter D. Ashlock; Universi-
dad Central de Venezuela, Instituto de Zoolo-
gia Agricola, Maracay, the late Dr. F. J. Fer-
nandez- Yepes.

Literature Cited

epilations prior to 195(i arc cited in Metcalf (1964).

Kramer, J. P. 1963. New Neotropical Neobalinae with
keys to the genera and to the .species of Conala
(Homoptera: Cicadellidae). Proceedings of the
Entomological Society of Washington 65: 201-
210.

LiNNAVUORi, R. 1959. Revision of the Neotropical Delto-
cephalinae and some related subfamilies (Ho-
moptera). Annales Zoologici Societatis Zoologicae
Botanicae Fennicae 'Vanamo' 20(1); 1-369.

LiNNAVUORi, R , AND F Heller 1961. Beitrag zur Ci-
cadelliden, Fauna von Peru. Stuttgarter Beitrage
zur Naturkunde 67: 1-14.

Metcalf, Z. P 1964. General catalogue of the Ho-
moptera, Fascicle VI, Cicadelloidea, bibliography
of the Cicadelloidea. Agricultural Research Ser-
vice, United States Department of Agriculture.
349 pp.

1966. General catalogue of the Homoptera, Fasci-
cle VI, Cicadelloidea, Part 16, Idioceridae. Agri-
cultural Research Service, United States Depart-
ment of Agriculture. 237 pp.

1967. General catalogue of the Homoptera, Fasci-
cle VI, Cicadelloidea, Part 10, Euscelidae. Agri-
cultural Research Service, United States Depart-
ment of Agriculture. 2,695 pp.

A NEW SPECIES OF PAZU OMAN FROM EASTERN CALIFORNIA
(HOMOPTERA: CICADELLIDAE: DELTOCEPHALINAE)

Raymond J. Gill'

Abstract — A new species of Pazu, P. monoinyo, from the Inyo-Mono County area of California is described.
Additional occurrence records are provided for Pazu halli (Beamer), and some of its morphological characteristics are
illustrated and compared with P. monoinyo.

The monobasic genus Pazu was erected by
Oman (1949) to include Hebecephalus halli
Beamer (1946) from central Arizona. Pazu is
distinguished from Hebecephalus primarily
by the shape of the styles, which in Hebe-
cephalus have the apical portion bent laterad
to the shaft at right angles, forming an elon-
gate, slender "toe." The type species oiPazu
and the new species described here have
more conventional, straighter styles (see Figs.
IE and 2E).

Specimens of this new taxon were collected
from antifreeze (ethylene glycol) pitfall traps
placed in sandy areas or on small sand dune
habitats in a 78-km" area in Inyo and Mono
counties near Bishop, California, and from a
single location 42 km east of Bishop near Deep
Springs. The area is semiarid Great Basin
habitat irrigated in the lower areas by
snowmelt runoff from nearby mountain peaks
or from artesian springs. The traps were
placed at these and other sand dune locations
throughout the southwestern U.S. as part of a
joint project between the California Depart-
ment of Food and Agriculture and the
U.S. Bureau of Land Management (BLM)
(Andrews et al. 1979). The purpose of the
project was to study the insect fauna associ-
ated with sand dune habitats as part of an
effort by the BLM to preserve sand dune habi-
tats.

The fact that all known specimens of P.
monoinyo exhibit microptery and have been
collected from just three pitfall trap locations
indicates that the species may spend consider-
able time on the soil surface and on the crowns
of the host rather than on aerial plant parts.
Whether a fully macropterous form capable of

flight actually exists in certain situations is
unknown. A macropter collected at Blythe,
Riverside County, California, on 2-IV-63 is
nearly identical in color pattern to P. mono-
imjo; however, the specimen is missing the
abdomen, thus making accurate identification
impossible. If such a form does exist, it is
probably not common and may explain why
the species has previously gone undetected.
In all collection locations the dry sand dune
habitats were in close proximity to water and
the lush vegetative growth associated with it.
The dune site east of Bishop, California, is
within 100 m of the Owens River; the Fish
Slough and Deep Springs locations are adja-
cent to a series of artesian springs. Since the
species has not been collected from other sand
dune localities, it would appear that the spe-
cies lives and feeds in or on the crowns of
plants, presumably grasses, near permanent
sources of surface water. However, several
attempts to re-collect the species at two of the
known localities have failed.

Pazu monoinyo, n. sp.

Description. — Resembles P. balli (Beamer)
but with darker, more distinct markings and
different genitalia.

External CHARACTERISTICS. — Groundcolor
apparently cinereous, although the possible
color-altering effects of the antifreeze and al-
cohol preservatives leave the actual color in
doubt. Markings fuscous. Crown with a
broad, longitudinal pair of lines, each with a
central cinereous line, so that the markings
appear as two pairs of fuscous lines on either
side of the midline; with a fuscous spot be-
tween each ocellus and compound eye. Frons

Analysis and IdentiPication, Division of Plant Industry, California Department of FcKid and Agriculture. 1220 N Street, Sacramento, California 95814.

1988

GiLL: A New Pazu from California

Fig. 1. Pazu rnonoimjo, n. sp.: A, adult female habitus; B, male aedeagus and connective, dorsal view; C, male
aedeagus and connective, lateral view; D, apex of male abdomen, lateral view; E, apex of male abdomen, ventral view.

with two pairs of arclike, subapical, fuscous
markings and occasionally with a pair of
rectangular, basal, fuscous spots next to
clypeus; frons apparently never clouded with
fuscous as in P. balli. Thorax with six fuscous,
longitudinal lines, thickest lines medial, all
lines lighter than those on crown. Abdominal
dorsum with three pairs of longitudinal, fus-
cous lines, the lateral pairs often coalescing to
form bands twice the width of medial pair.
Venter cinereous variously marked with fus-
cous, especially on anterior median areas of
sternites. Legs cinereous with two or more
fuscous spots or bars on femora, other leg
segments darker near spines and at spine
bases. Elytra reduced (micropterous as delin-
eated by Oman 1987), barely reaching ninth
abdominal segment. Most elytral cells filled to
a greater or lesser degree with fuscous; all
veins or cells on either side of claval vein
usually clear or white. Wings reduced, ca 1/2 x

length of elytra. There appears to be no color
or other dimorphisms between the sexes.

Males — Length 3.4-3.7 mm (mean 3.6
mm), specimens slightly shriveled due to
preservation methods. Wing length 2.4-2.7
mm (mean 2.6 mm); vertex length 0.4-0.6
mm (mean 0.5 mm); transocular width
1.0-1.3 mm (mean 1.2 mm); interocular
width 0.4-0.6 mm (mean 0.6 mm). Pygofer
shorter than in P. balli, more truncated, with
just a vestige of a tooth on lateroventral cor-
ners; weakly setose. Plates evenly tapered to
acute apices and completely divided, not
basally fused and truncate as in P. balli; spine-
like setae uniseriate. Style apices J-shaped,
medial apex long, slender, very slightly
curved laterad; lateral tooth broadly triangu-
lar. Connective linear, about as long as aedea-
gal shaft. Aedeagus roughly V-shaped; ante-
rior arm broader, apex curved posteriorly;

Great Basin Naturalist Memoirs

No. 12

Fig. 2. Pazu halli (Beamer): A, adult female habitus; B, male aedeagu.s and connective, dorsal view; C, male aedeagus
and connective, lateral view; D, ape.x of male abdomen, ventral view.

posterior arm ca 1/2 x width of anterior, gradu-
ally and evenly tapered to apex, crescentic in
lateral view. Aedeagal apex with three pairs of
apicolateral flanges or blunt teeth.

Females. — Length 4.6-5.0 mm (mean 4.6
mm). Wing length 2.9-3.1 mm (mean 3.0
mm); vertex length 0.7-0.9 mm (mean 0.7
mm); transocular width 1.4-1.6 mm (mean
1.4 mm); interoeular width 0.6-0.9 mm
(mean 0.6 mm). Last ventral segment with
broadly rounded, posteriorly directed median
lobe.

Etymology. — The name monoinyo is a
noun in apposition referring to the two locali-
ties where the species was collected. Mono
and Inyo counties, California.

Discussion. — Pazu monoinyo is similar to
the generic type, P. halli, in color pattern and
male genitalia. Color patterns differ in that
the dorsal, longitudinal lines tend to be much
darker and more distinct on the head and
thorax in P. monoinyo, but much lighter and
with a tendencv to coalesce in P. halli. The

male genitalia differ in that the pygofer is
much shorter in P. monoinyo; the plates are
completely divided and have acute apices in
P. rnonoinyo compared with basally fused,
truncate plates in P. halli. The last ventral
segment of the female is lobed posteriorly in
P. monoinyo but deeply cleft in P. halli. All
known specimens of P. monoinyo exhibit mi-
croptery, whereas all those of P. halli are
macropterous.

Specimens of P. halli are known from
Cochise and Pearce, Cochise County, Ari-
zona.

Material examined. — Male holotype and
female allotype: "Calif, Mono Co., 9 mi N
Bishop, Fish Slough, 4,200', sand dunes, V-
12-82 to XII-20-82, Derham Giuliani, col-
lected in ethylene glycol pit trap." Paratypes:
"Calif, Mono Co., 9 mi N Bishop, Fish
Slough, 4,200', .sand dunes, V-12-82 to XII-
20-82, Derham Giuliani, collected in ethyl-
ene glycol pit trap," 4 males and 2 females;
"Calif, 3 mi E Bishop, 4,100', sand dunes,

1988

GiLL: A New Pazu from California

V- 12-82 to XII-10-82, antifreeze pit trap, D.
Giuliani," 1 male and 4 females; "Inyo Go.,
Ga. , Deep Springs Valley, VI-9-78, antifreeze
pit trap, D. Giuliani," 1 female.

Type deposition. — The holotype and allo-
type (both GAS Type # 16039) were deposited
in the Galifornia Academy of Sciences, San
Francisco (GAS). Twelve paratypes were de-
posited as follows: 2 males, 2 females, San
Francisco (GAS); 2 males, 4 females, Galifor-
nia Department of Food and Agriculture,
Sacramento (GDFA); 1 male, 1 female.
United States National Museum, Washing-
ton, D.G. (USNM).

Acknowledgments

Thanks must go first to Dr. Paul Oman, who
has been such a great source of inspiration for
all of us who work on the Gicadellidae. His
publications have been extremely valuable to
me in the course of my work as a taxonomist
and identifier, and his 1949 book on the
Nearctic lealhoppers was the primary build-
ing block toward my understanding the Gi-
cadellidae.

I express gratitude to Dr. Leon W. Hep-
ner, Mississippi State University, State Gol-

lege, Mississippi, and to Dr. Steve Wilson,
Gentral Missouri State University, Warrens-
burg, Missouri, for reading the manuscript
and offering useful suggestions. I also thank
Dr. Robert Brooks, Snow Entomological Mu-
seum, Lawrence, Kansas, for the loan of the
type specimens oi Pazu balli. I am extremely
grateful to Susan Sawyer, GDFA, for the
habitus sketches of Pazu monoinyo and Pazu
balli.

Literature Gited

Andrews, F G , A R Hardy, and D Giuliani 1979. The
coleopterous fauna of selected California sand
dunes. Report to the ELM on contract CA-960-
128.5-1288 DEOO, California Department of
Food and Agriculture, Sacramento. 142 pp.

Reamer, R. H 1936. New leafhoppers from the western
United States (Homoptera: Gicadellidae). Cana-
dian Entomol. 68: 2.52-2.57.

Oman, P W 1949. The Nearctic leafhoppers (Ho-
moptera: Gicadellidae). A generic classification
and checklist. Washington Entomol. Soc. Mem. 3:
1-2.53.

1987. Alary polymorphism in the Gicadellidae and

its ecological implications. Pages 55-63 in M. R.
Wilson and L. R. Nault, eds., Proc. 2nd Int.
Workshop on Leafhoppers and Planthoppers of
Economic Importance, Provo, Utah, USA, 28
July-1 August 1986. GIE, London.

EVIDENCE FOR AN INDO-PACIFIC ORIGIN OF HAWAIIAN ENDEMICS
IN BALCLUTHA AND RELATED GENERA (CICADELLIDAE: MACROSTELINI)

W. J. Knight' and M. D. Webb'

Abstract — The 10 endemic species of Balclittha in Hawaii are reviewed and compared with the Pacific and
American forms of the genus. Two new species, nigriventris and fiiscifrons , are described; kilaueae is transferred to a
new genus Balolina n. comb. ; beardsleyi is synonymized with saltuella , n. syn. ; and hospes is relegated to a subspecies
oiincisa, n. stat. Descriptions and notes, together with illustrations and a key, are given for each taxon. Evidence is
presented to suggest that all the endemic species oi Balclittha have arisen within the islands from a single colonization
and that the taxon showing the greatest similarity to them is the widespread Pacific species B. lucida. It is shown that
the Hawaiian genus Nesolina is most closely related to the endemic species-group oi Balclutha and that Balolina is an
independent colonization.

The isolated position, volcanic origin, and
diverse habitats of Hawaii, together with the
division of the area into several large islands,
have combined to produce a depauperate, yet
large and highly endemic, biota (Simon 1987).
This phenomenon is well illustrated by the
plant-feeding insect family Cicadellidae. The
family was studied in the islands by Kirkaldy
(1910) and Osborn (1935) and subsequently
reviewed by Zimmerman (1948). Namba
(1956) later described several new species.

In his review of the Hawaiian insect fauna,
Zimmerman (1948) considered the generic
affinities to be predominantly Pacific and only
5% American. The Homoptera were consid-
ered to be exclusively Pacific, although little
evidence was provided.

Compared with the Pacific and mainland
America, the Hawaiian cicadellid fauna is very
depauperate at the supraspecific level. Only 5
of the 35 subfamilies are present, 4 of which
are represented by 5 recently introduced spe-
cies from North America. In contrast, the
Deltocephalinae, a large, worldwide group
with 17 currently recognized tribes, is repre-
sented by 2 tribes. The endemic genera
Nesophryne (2 species), Kirkaldiella (2 spe-
cies), and Nesophrosyne (60 species) belong to
the Opsiini and are related to the Pacific
genus Orosius (Linnavuori 1960, Ghauri
1966). Nesolina (1 species), Balolina n. gen. (1
species), and the cosmopolitan genus Bal-
clutha, comprising 105 species and repre-
sented in Hawaii by 11 endemic and 2 cos-

mopolitan species, belong to the Macros-
telini, the major cicadellid group in the simi-
larly isolated islands of Marquesas in the
Pacific, and St. Helena in the Atlantic. This
tribe contains approximately 30 genera world-
wide, of which 10 are island endemics, 2 in
Hawaii, 4 in the Marquesas, and 4 on St.
Helena. Interestingly, although the main host
plants of the group are Gramineae, in Hawaii,
Marquesas, and St. Helena shrubs and tree
ferns of southern temperate distribution are
also utilized (Webb 1987).

Materials

The material studied is primarily from the
Bishop Museum, together with some in the
British Museum (Natural History), abbrevi-
ated throughout the text to BPBM and
BMNH, respectively.

Methodology

The present study investigates the origins
of the Hawaiian Balclutha species. This has
necessitated a reexamination of all the Hawai-
ian species and a review of the variation in
characters among Pacific and American spe-
cies of the genus. Until a phylogenetic analy-
sis of the genus and tribe is undertaken, any
statements on the polarity of characters within
these taxa would be premature. Our bio-
geographical inquiry is therefore based on
phenetic resemblances rather than on shared

British Museum (Natural History), Cromwell Road, London SW7 5BD, Enfiland.

1988

Knight, Webb: Hawaiian Endemics in Balclutha

apomorphies and should be seen as only ex-
ploratory and preliminary in nature.

Faunal Relationships of
Hawaiian Balclutha

The variation in morphological characters
in Balclutha was reviewed by Knight (1987)
for the south and west Pacific species and by
Blocker (1967) for the American species. Both
faunas are of comparable size, 30 and 32 spe-
cies, respectively, but have only 3 species in
common, saltuella, incisa, and lucida.

Compared with the Pacific and American
species, the 10 endemic Hawaiian species,
nigriventris n. sp., fuscifrons n. sp., grandis
Namba, phoxocephala Namba, timberlakei
(Osborn), lohata Namba, usitata Namba, vol-
canicola (Kirkaldy), plutonis (Kirkaldy), and
peregrina (Kirkaldy), differ only slightly from
each other in head shape and minor details X)f
the male genitalia, suggesting that they prob-
ably arose from one founder species. The
character states common to the 10 endemic
species are listed below, the condition
present in the 10 endemic species shown in
italics:

1. Relative width of head and pronotum: head equal to
or wider than the pronotum.

2. Length of vertex: longer medially than next to eyes.

3. Laterofrontal sutures: terminating at or near the
ocelli.

4. Distance between ocelli and eyes: ocelli separated
from the eyes by a distance equal to 2-4 times their
own diameter.

5. Foretibia setal formula; 1.3.

6. Hind femora setal formula: 2. i.i.

7. Shape of male pygofer: simple, without ventral lobes
or processes.

8. Shape of subgenital plates: triangtdate, with apical,
fingerlike process 1/4-1/2 total length of plate.

9. Shape of styles: robust, with apical process and
preapical lobe well developed.

10. Relative length of stem and arms of connective: stem
longer than arms.

11. Shape of aedeagal shaft: simple, elongate, curving
dorsally or ante r odor sally to point level with or
slightly anterior to atrium.

Comparison between the Hawaiian,
Pacific, and American Species

In the present section, the Pacific and
American species of the genus are reviewed
for each of the characters listed above. The
alternative conditions of characters 3, 5, and
7 are coded A and B for ease of reference in

subsequent discussion.

The relative width of the head (character 1)
varies within the genus and may be wider,
equal to, or narrower than that of the prono-
tum. All three conditions are present in both
the American and the Pacific faunas, half the
species in each region showing the head ap-
proximately equal to or wider than the prono-
tum, as in Hawaii.

The vertex (character 2) is usually short and
uniform in length and is one of the main dis-
tinguishing features of the genus. The medial
prolongation of the vertex found in the Hawai-
ian species occurs in the Pacific in only
smaragdula , and in a few individuals offlagel-
lata, bifasciata, rieki, and the cosmopolitan
species punctata. It is far more prevalent
among the American forms, being a character-
istic feature of 7 species and occurring occa-
sionally in 15 others.

The laterofrontal sutures (character 3) usu-
ally terminate at or near the ocelli (condition
B) but are occasionally turned medially, ven-
trad of the ocelli (condition A). With the ex-
ception of simplex and the cosmopolitan
saltuella , the Pacific species have the sutures
terminating near the ocelli as in Hawaiian
species. Both conditions also occur in the
American fauna but with an approximately
equal number of species of each.

The ocelli (character 4) are usually small
and situated on the anterior margin of the
head, separated from the corresponding eye
by approximately their own diameter. Most of
the American and Pacific species have the
ocelli in this position, or only slightly more
remote from the eye. Only i7npicta and arc-
tica in America, flagellata and cheesmanae in
the Pacific, and the cosmopolitan species
punctata, have the ocelli more than twice
their own diameter from the eyes, and ap-
proximating the condition found in Hawaiian
species. The Pacific fauna shows greater flexi-
bility in this character, however, with chlor-
optera and hulhosa having the ocelli enlarged
and more dorsad in position.

Throughout the genus, the foretibia setal
formula (character 5) may be 1. 1 (condition A),
1.3 (condition B), or rarely 1.4. The condition
in Hawaii (condition B) is similar to that in the
majority of Pacific species. Only two Pacific
species, simplex and the cosmopolitan saltu-
ella, show condition A. In the American
fauna, half of the species examined have con-
dition A and half condition B.

Great Basin Naturalist Memoirs

No. 12

The setal pattern at the apex of the hind
femora (character 6) varies within the genus
from 2. 1. 1 to 2.2. 1. The two conditions occur
equally throughout both the Pacific and
American faunas, unlike Hawaii where only
the former is found.

The male pygofer (character 7) varies from a
simple, rounded shape (condition A), as found
in Hawaii, to a more complex form with pos-
teroventral lobes or processes (condition B).
The simple form is rare outside Hawaii, occur-
ring in only simplex in the Pacific and in the
species lucida and saltuella, which occur in
both regions.

The subgenital plates (character 8) are char-
acterized in all species by a membranous, fin-
gerlike, apical segment. The shape of the
plate found in the Hawaiian species occurs in
the majority of Pacific and American species,
being slightly more prevalent in the latter
region. The remaining species in both regions
have the basal part of the plate much reduced
in size.

The robust, well-developed styles (charac-
ter 9) of the Hawaiian species occur in the
majority of species in both the Pacific and
America but are more widespread in the for-
mer. Only two Pacific species, bulbosa and
rieki, have a reduced style, compared with
nine species in the Americas, plus the cosmo-
politan saltuella in both regions.

The Y-shaped connective (character 10)
varies principally in the relative length of the
stem and arms. The condition present in
Hawaii, with the stem longer than the arms, is
present in 50% of the species in the Pacific but
in only 30% of the species in America.

A simple, filamentous aedeagus (character
11) is characteristic of the majority of species
in the genus. In the Hawaiian species the shaft
is directed dorsally or curved anterodorsally
to a point approximately level with the well-
developed, basal apodeme, a form that occurs
in 50% of the fauna in both the Pacific and
America.

Faunal Similarity

If we take the number of species in each
region showing character states identical with
those in Hawaii as a measure of similarity,
then the Hawaiian species show greater simi-
larity to the Pacific fauna with respect to the
laterofrontal sutures, foretibial setal pattern,
styles, connective, and to a lesser extent the

ocelli and male pygofer. The American spe-
cies show greater similarity only in the length
of the vertex and to a lesser extent in the shape
of the subgenital plates. The relative width of
the head, the setal pattern on the hind
femora, and the shape of the aedeagus provide
little information in this respect. If we elimi-
nate those characters that confer only minimal
regional preference, such as the ocelli, male
pygofer, and subgenital plates, and also those
known to vary infraspecifically, such as the
length of the vertex, we are left with only four,
the laterofrontal sutures, the foretibial setal
pattern, and the shape of the styles and con-
nective, all of which support the theory of
closer links with the Pacific.

Further support for a Pacific link is pro-
vided by the Sorensen Coefficient of Similar-
ity (Southwood 1978)

Cs = 2j/(a + b)
where j is the number of character states com-
mon to the two areas and a -I- b are respec-
tively the number of character states in each
area. Based on the endemic species in each
area, the coefficient for Hawaii and the Pacific
is 0.74 and for Hawaii and the Americas is
0.58, showing a greater degree of similarity
between the first two areas.

From the above review it is seen that no
one character is shared exclusively between
Hawaii and either of the other two regions to
provide evidence of affinity. However, if we
consider a combination of characters, rather
than single characters alone, then the picture
becomes a little clearer.

Table 1 shows the Pacific species and their
similarity or dissimilarity to the Hawaiian en-
demic species with regard to the 11 charac-
ters. This reveals an obvious correlation be-
tween characters 3, 5, and 7. All three
characters show condition A" in the first two
species and condition B" in the fointh species
onward. An intermediate stage occurs in the
third species, lucida , which has condition B in
characters 3 and 5 but retains condition A in
character 7. The American species show the
same correlation between characters 3, 5, and
7, but, except for lucida , the character states
in the intermediate stage are reversed. This is
shown in Table 2, where the intermediate
stage is highlighted for each region. The simi-
larity between the Hawaiian endemics and
lucida in this respect is unique.

; scttion, paragraphs 4, 6. ami H.

1988

Knight, Webb: Hawaiian Endemics in Balclutha

Table 1. A list of the Pacific species oi Balcltttha. showinjz; the similarities ( + ) and differences ( — ) between them and
the Hawaiian endemics for the 1 1 listed characters. (For further explanation, see text.)

Pacific

Characters

species

saltuella

—

simplex

lucida

—

bilobata

batuensis

punctata

neglecta

viridinervis

flageUata

bifasciata

■±

trilineata

rieki

■±_

cheesinanae

yanchepensis

-H

smaragdula

alstoni

kuroiwae

asyrmnetrica

chloroptera

bulbosa

bacchusi

spiniloba

incisa

rosea

pseudorosea

rubrostriata

wilsoni

distenda

riibrinervis

noonadana

Table 2. Character-state distribution for three characters in Pacific and American species of Balclutha. (For further
explanation, see text.)

Characters

American
species*

Chi

iracters

species

saltuella

simplex

neglecta

lucida

chiasma
sandersi

A
A

B
B

remaining spp.

*lucida. which occurs in Central and South America, has been omitted from this half of the table in the interests of clarity.

Distribution of B. lucida

Since lucida is the species most similar to
the Hawaiian forms, it is of interest to examine
its distribution relative to Hawaii. Figure 53
shows its distribution, extending from the
Aldabra Islands (Webb 1980 \filum]) in the
west to Central and South America in the east
(Blocker 1967 \jloridana ] and Knight 1987). It
is a tropical and semitropical species clearly

capable of movement over long distances and
with the ability to colonize and establish itself
on islands and mainland areas. Its presence in
the Marshall and Marquesas islands places it
within a comparatively short distance from
Hawaii. The available data, however, provide
no information on the center of origin for the
species, although its absence from mainland
Africa and Asia suggests that it is either the
Pacific or Central America.

Great Basin Naturalist Memoirs

No. 12

Relationship of Balclutha to Other
Hawaiian Macrostelini

In addition to Balclutha , two other macros-
teline genera occur in Hawaii, both endemic.

The monotypic genus Nesolina differs from
Balclutha principally in the more elongate
head and the possession of longitudinal brown
stripes on the vertex and pronotum. In all
other characters it closely resembles the en-
demic species of Balclutha to which it is prob-
ably most closely related. Despite this close
similarity, the present status of Nesolina is
preserved, pending a phylogenetic study of
the Macrostelini.

The relationship of the remaining Hawaiian
macrosteline genus, Balolina n. gen., is un-
certain. Although it is similar in general ap-
pearance to Balclutha and the Pacific genus
Nesoclutha and has the first valvulae fused as
in Balclutha (Fig. 33), it differs in many signif-
icant features, as listed below in the key and
generic description. Except for the position of
the ocelli, there is little to indicate that it is in
any way related to the Hawaiian species of
Balclutha . On the contrary, it appears to be
an independent introduction to the islands.

All three Hawaiian macrosteline genera be-
long to a group having the hindwing veins sc
and r united into a single vein, with a conse-
quent reduction in the number of apical cells
(Fig. 32). This is considered by Ossiannilsson
(1983) to be justification for placing Balclutha
in a separate tribe, the Balcluthini, which is
also characterized (Oman 1949) by the posses-
sion of a sulcate hind basal tarsus (Fig. 74).
Both characters are found to some degree,
however, in other genera, including Macro-
steles . The phylogeny of the Macrostelini pro-
posed by Triplehorn and Nault (1985) in-
cludes Balclutha but omits several other gen-
era, including the 10 island endemics. Some
of these have been treated by Webb (1986,
1987).

Distribution of Hawaiian Macrostelini

Table 3 summarizes previous (x) and new
(N) distribution data and shows that the
largest and youngest islands, Hawaii and
Maui, respectively, support the most species,
and that Kauai and Molokai, which differ con-
siderably in size, have similar numbers of spe-
cies. Further collecting on the islands may

have profound effects on the distributional
records and may even reveal further species.

Keys and Descriptions

The taxonomy of the Hawaiian Macrostelini
has been treated by several authors (see in-
troduction), but descriptions are usually
incomplete and contain little information on
intraspecific differences. Namba's (1956) revi-
sion o( Balclutha was a considerable improve-
ment on previous work, but a few of the char-
acters referred to are unreliable. New keys to
genera and species are given to take account of
this and to accommodate the new taxa. All
host records given below are taken from ei-
ther Zimmerman (1948), Namba (1956), or
specimens examined.

Kev to the Genera of Hawaiian Macrostelini

1. Head and thorax with longitudinal, brown

bands; vertex elongate (Fig. 54) Nesolina

— Head and thorax not marked as above; vertex
short to moderately long (Figs. 7, 15) 2

2(1). Vertex moderately long, ocelli situated almost
to point midway between eye and midlength of
foremargin of vertex (Fig. 67). Clypellus ex-
tending well beyond margin of face, trans-
clypeal suture obscure; laterofrontal sutures
short (Fig. 68). Foretibia with apical setae simi-
lar in length (Fig. 73). Hind femur with apical
setal formula 2 + 2+1 (Fig. 69). Macrosetae of
legs and genital segment nonhairy. Aedeagal
shaft with pair of apical processes (Fig. 82);
gonopore apical on ventral surface (Fig. 76).
Connective with stem very short (Fig. 80).
Subgenital plate with irregular macrosetae
(Fig. 83). Second valvulae evenly tapered to
apex; teeth extending over distal half of dorsal
margin and onto apex of ventral margin (Fig.
75) Balolina

— Vertex short to moderately long; ocelli situated
near eye or at point one-third distance from eye
to midlength of foremargin of vertex (Figs. 2,
15, 28). Foretibia with apical setae dissimilar in
length (Fig. .35). Hind femur with apical setal
formula 2+1 + 1 (Fig. 30). Male and female
genitalia not as above Balclutha

Key to Species of Hawaiian Balclutha

1. Vertex of uniform length; ocelli 1.0-1.5 times
own diameter from corresponding eye; coronal
suture visible in facial view (Figs. 28, 29)
Pygofer lobe with a ventral process (Figs. 37,
46). Connective with arms ecjual in length to
stem (Fig. 42) 11

— Not as above 2

2(1). Yellow to greenish yellow, marked with brown
on first thoracic sternite (as in saltuella. Fig.

1988

Knight, Webb: Hawaiian Endemics in Balclutha

Table 3. Distribution ofthe Macrostt'lini on the Hawaiian islands, from the oldest in the north (left) to the youngest
in the south (right).

Area (sq km)

Kauai

1437

Oahu
1564

Molokai

673

Lanai
365

Maui
1885

Hawaii
10437

Nesolina
line at a

Balolina
kilaueae

Balclutha
grandis
saltuella
phoxocephala
incisa hospes
timberlakei
lobata
usitata
volcanicola
plutonis
peregrina
nigriventris
fuscifrons

Total

N
N
10

43), abdomen (at least dorsally), and sometimes
legs 3

— Not marked as above or if abdomen and first
thoracic sternite marked with brown, then
overall color yellow tinged with scarlet 5

3(2). Large species, 4.0-4.4 mm in length. Aedea-

gus and style as in Figures 3, 4, 18

fuscifrons. n. sp.

— Small to moderately large species, up to 3.5
mm 4

4(3). Vertex marked with stramineous or brown
(Figs. 5, 7). Aedeagal shaft robust (Fig. 24) . . .
volcanicola (Kirkaldy)

— Vertex without markings. Aedeagal shaft slen-
der (Fig. 19) nigriventris, n. sp.

5(2). Large species 4.5-5.3 mm in length. Stem of
connective broad. Aedeagus as in Figure 21 . .
grandis Namba

— Small to moderately large species, up to 4.0
mm 6

6(5). Preapical lobe of style large; apical process
abruptly tapered distally (Figs. 16, 17). Female
genital sternite marked with brown on disc

(Fig. 8) lobata Namba

— Not as above 7

7(6). Moderately large species, 3.5-4.0 mm.
Clypellus with sides concave. Connective
elongate, longer than style (as in timberlakei.
Figs. 13, 14). Aedeagal shaft slender, evenly
curved dorsally (Figs. 25, 26) . plutonis (Kirkaldy)

— Small species, 2.6-3.1 mm; without the above
combination of characters 8

8(7). Aedeagal shaft slender (Figs. 25, 27) 9

— Aedeagal shaft robust (Figs. 22, 23) 10

9(8). Clypellus with sides straight (Fig. 12). Aedea-
gal shaft evenly curved throughout length (as in

plutonis. Fig. 25) peregrina (Kirkaldy)

— Clypellus with sides concave. Aedeagal shaft

angularly curved basally (Fig. 27)

usitata Namba

10(8). Aedeagal shaft sinuate (Fig. 22). Connective
elongate, longer than style (Figs. 13, 14) ....
timberlakei (Osborn)

— Aedeagal shaft not sinuate (Fig. 23). Connec-
tive approximately equal in length to style . . .
phoxocephala Namba

11(1). Female pregenital sternite with disc marked
with brown (Fig. 36). Male genitalia as in Fig-
ures 37-40 incisa hospes (Kirkaldy)

— Female pregenital sternite not marked with
brown. Male genitalia as in Figures 41, 42, 45,

46 saltuella (Kirschbaum)

Balclutha Kirkaldy

Balclutha Kirkaldy, 1900: 243. New name for Gnathodus
Fieber, 1866; Blocker 1967; 4, Knight 1987: 1178.
Type species: Cicada punctata Fabricius, 1775,
by monotypy.

Balclutha fuscifrons , n. sp.
Figs. 1-4, 18

Length. — S , 4.0-4.2 mm; 9 , 4.2-4.4 mm.

Yellow tinged with green, abdomen heavily
marked dorsally and ventrally with dark
brown; clypeus pale to dark brown; legs and
thoracic sternites variably marked with dark
brown.

Head approximately equal in width to
pronotum. Vertex with medial length approx-
imately 1.33 times length next to eye. Ocelli
marginal, separated from corresponding eye
by distance equal to 3-4 times own diameter.

Great Basin Naturalist Memoirs

No. 12

Figs. 1-17. Endemic Hawaiian Balclutlia species. 1-4, B. fuscifrons: 1, male pygofer, left lateral view; 2, head and
thorax, dorsal view; 3, right style, dorsal view; 4, connective, dorsal view. 5-7, B. volcanicohi: 5. head and thorax,
dorsal view (Kilauea); 6, right style, dorsal view; 7, head and th()ra.\, dorsal view (Wailua). 8, B. lohata. female
pregenital sternite. 9-11, B. nigriventris: 9, right style, dorsal view; 10, connective, dorsal view; 11, head and thorax,
dorsal view. 12, B. peregrina, face. 13-15, B. timhcrlakci: 13, right style, dorsal view; 14, connective, dorsal view; 15,
head and thorax, dorsal view. 16-17, B. lohata, apex of right style, dorsal view.

Face slightly wider than long; clypellu.s with
sides concave, apex extending very slightly
beyond margin of face; laterofrontal sutures

convex, distally concave, reaching to corre-
sponding ocellus. Setal formulae: foretibia,
1.3 or 1.4; midtibia, 4.3, 4.4, 5.3, or 5.4; hind

1988

Knight, Webb: Hawaiian Endemics in Balclutha

femur, 2.1.1 or 2.1.1.1.

Male. — Pygofer simple, posterior margin
broadly and evenly rounded in lateral aspect.
Subgenital plate tapered distally to moder-
ately long, fingerlike process. Connective
with arms shorter than stem. Style approxi-
mately equal in length to connective, preapi-
cal lobe elongate. Aedeagal shaft slender,
more or less straight distally.

Female. — Pregenital sternite with poste-
rior margin more or less straight. Valvulae as
in incisa (Figs. 33, 38).

Host. — Unknown.

Distribution. — Endemic (Maui).

HOLOTYPE. — 6, Maui, Kula Pipe Line,
4,200', vii.1956, R. Namba (BPBM).
Paratypes: Maui: 13(5, 199, same data as
holotype; 4c5, Holua, Haleakala Crater,
vi.l983, D. E. Hardy; 4c?, 99, Waikamoi,
4,000', D.E. Hardy (BPBM, BMNH).

Remarks. — This species can be distin-
guished by its large size, brown markings, and
more or less straight aedeagal shaft. Its exter-
nal appearance is similar to Balolina hilaueae,
with which it may be confused.

Balclutha volcanicola (Kirkaldy)

Figs. 5-7, 24

Nesosteles volcanicola Kirkaldy, 1910; 574: Osborn 1935;

58, Fig. 24e. Lectotype S, Hawaii (BPBM), here

designated [examined].
Balclutha volcanicola (Kirkaldv); Zimmerman 1948: 87,

Fig. 27, Namba 1956; 107-108, Fig. 7.

Host. — Cyperus (Cyperaceae), Eragrostis
(Gramineae), Lythrum (Lythraceae).

Distribution. — Endemic (Molokai, Maui,
Hawaii).

Material examined. — Lectotype <5, Ha-
waii, Kilauea. Maui: \6 , Wailua (BPBM).
Hawaii: Id, paralectotype, same data as lec-
totype, M, 1 abdomen missing, Kilauea, on
Lyf/inm (BPBM); 4c?, Kilauea (BMNH); 4(5,
Kealakekua(BMNH).

Remarks. — This species can be distin-
guished by its small size (c?, 2.6 mm, 9 , 2.8
mm), stramineous or brown markings on the
head (Figs. 5, 7), brown markings on the first
thoracic sternite and dorsum of the abdomen,
the head being distinctly wider than the
pronotum (Figs. 5, 7), 3.3 midtibial setal for-
mula, robust aedeagal shaft (Fig. 24), and the
moderately long preapical lobe on the style
(Fig. 6).

The medial length of the vertex varies from

slightly longer than to 1.5 times length next to
eyes (Figs. 5, 7).

Balclutha nigriventris , n. sp.
Figs. 9-11, 19

Length.— (5, 3.0-3.3 mm; 9, 3.2-3.5 mm.

Yellow to greenish yellow, variably tinged
with orange to brown on head, venter, and
legs. Dorsum of abdomen dark brown.

External characters as in fuscifrons .

Male. — Genitalia as in fuscifrons but with
aedeagal shaft curved slightly more dorsally
(Fig. 19).

Female. — Genitalia with posterior margin
of pregenital sternite more or less straight.
Valvulae as in incisa (Figs. 33, 38).

Host. — Deschampsia (Gramineae).

Distribution. — Endemic (Maui, Hawaii).

Holotype. — S, Hawaii: Hualalai, 6,000-
7,000', 21. iv. 1944, N. L. H. Krauss (BPBM).
Paratypes: Maui: 36, 39, Halemauu Trail,
1.V.1945, 8,000' (1(5, on Deschampsia), 16,
Holua, Haleakala Crater, 6,500', vi.l953.
Hawaii: 16, 19, same data as holotype;
19, Hualalai, Hinatapoula, 5,900', vii.1964
(BPBM, BMNH).

Remarks. — This species is similar to fus-
cifrons but is significantly smaller, often lacks
the brown marking on the clypellus, and has
the shaft of the aedeagus slightly different in
shape (compare Figs. 18, 19).

Balclutha grandis Namba

Fig. 21

Balclutha grandis Namba, 1956: 105-106, Figs. 4a-b.
Holotype cJ, Kauai (BPBM) [examined].

Host. — Unknown.

Distribution. — Endemic (Kauai).

Material examined. — Holotype 6 , Kauai:
Alakai Swamp, viii.1953, 4,000' (BPBM).

Remarks. — This species can be distin-
guished by its large size (6 , 4.5 mm, 9, 5.3
mm), the head being distinctly wider than the
pronotum, 4.4 midtibial setal formula, broad
stem of the connective, and the shape of the
aedeagus (Fig. 21). Namba (1956) recorded
1 c5 , 1 9 from the type locality.

Balclutha lobata Namba

Figs. 8, 16, 17, 20

Balclutha lobata Namba, 1956; 106, Figs. 5a-b. Holotype
6, Kauai (BPBM) [examined].

Host. — Cyathodes (Gramineae), Styphelia
(Epacridaceae).

Great Basin Naturalist Memoirs

No. 12

Figs 18-27. Endemic Hawaiian Balclutha species, aedeagus, left lateral view, 18, B.fuscifrons; 19, B. ni^riventris;
20, B. lobata; 21, B. grandis; 22, B. timbcrlakei; 23, B. phoxocephala; 24, B. volcanicola; 25-26, B. plutoms; 27, b.

usitata.

Distribution. — Endemic (Kauai, Oahu,
Molokai, Maui, Hawaii).

Material examined. — Holotype S, Kauai,
Kokee, vii.1952, 3,600' (BPBM). Kauai: 8(5,

49, Pihea, Kaunuohua Ridge, 4,260'; 6c?,
19, Kokee, on Styphelia. Molokai: 16, Ko-
moku; 9(5, 69, above Waikolu V. 1,400m;
16, Kalaupapa Lookout; 26, Kewala Gulch,

1988

Knight, Webb: Hawaiian Endemics in Balclutha

3,500'. Maui: 19, Kula Pipe Line, 4, 22'
(BMNH, BPBM).

Remarks — This species can be distin-
guished by its moderately large size (6,
3.0-3.3 mm, 9, 3.3-3.4 mm), its dark ap-
pearance resulting from the dark brown dor-
sum of the abdomen and the greenish tinge of
the forewings, the 3.3 midtibial setal formula,
the disc of the female pregenital stemite
marked with brown (Fig. 8j, and the large
preapical lobe of the style (Fig. 16j. Some
specimens (Molokai) have the lobe of the style
narrower (Fig. 17). Contrar\- to \amba(1956).
the head is approximately equal in width to
the pronotum.

Balclutha plutonis (Kirkaldy)
Figs. 2.5, 26

Nesosteles plutonis Kirkaldy, 1910: 574; Osbom 19.3.5:
57-58, Fig. 24d. Lectotype 6 , Hawaii, here des-
ignated [e.xamined].

Balclutha plutonis ( Kirkaldy i: Zimmerman 1948: 87, Fig.
27, Namba 1956: 105. Fig. -3.

Host. — Coprosma (Rubiaceael Vaccinium
reticulatus (Ericaceae), Sophora iLegumi-
nosae), Argyroxiphium virescens, Raillardia
(Compositae), Deschampsia , Eragrostisi?)
(Gramineae), Vincentia (Cx-peraceae).

Distribution. — Endemic (Maui, Hawaii).

Material EX.\MIN ED — Lectotype 6, Ha-
waii, Kilauea, vii.1906. R. C. L. Perkins
(BPBM). Numerous specimens from Maui
(Haleakala, Puu Luau, Puu Xoauiau) and
Hawaii (Pohakuloa, Kilauea, Mauna Loa,
Hualalai)(BPBM, BMNH).

Re.MARKS. — This species can be distin-
guished by its moderately large size \ i . 3.5-
3.6 mm, 9, 3.6-4.0 mm), its uniform yellow
or greenish yellow color, usuall\- without
browTi markings on dorsum of abdomen, and
the elongate, evenly cur\ed shaft of the
aedeagus. It is similar to peregrina but is
larger, has the shaft of the aedeagus usually
slightly longer, the connective longer than
rather than equal in length to the styles, and
the sides of the clypellus conca% e rather than
straight. The lectot>pe is the only specimen
knowTi to have the dorsum of the abdomen
browTi and a slightly shorter aedeagal shaft
(Fig. 25). similar to peregrina . The setal for-
mulae var\- as follows: foretibia. 1.3. 1.4. or
1.5; midtibia, 4.4 or 5.4; hind femur. 2. 1. 1 or
2.1.1.1.

Balclutha peregrina (Kirkaldy)
Fig. 12

Nesosteles peregrina Kirkaldy, 1910: 575. Holotype 6,

Hawaii (BPBM) [examined].
Balclutha peregrina (Kirkaldy): Zimmerman 1948:

86-87, Fig. 27, Namba 19.56: 108, Figs. 8a-c.

Host — Cyathodes, Deschampsia (Gram-
ineae), Railliardia (Compositae), Vaccinium
(Ericaceae), Styphelia (Epacridaceae).

Distribution — Endemic (Maui, Hawaiij.

M.\TERIAL EX.^MINED — Holot>pe (5, Ha-
waii: Kilauea, vi.l903, R.C.L.P. Hawaii: (nu-
merous specimens from Kilaueaj; 26, 59,
Stainback Highwav, 2,134 m, on Styphelia
(BPBNL BMNH). '

Remarks — This species is similar to plu-
tonis but is significantly smaller (6 , 2.6-3.1
mm; 9 , 2.8-3. 1 mm ), has the abdomen some-
times marked with pale brown, the sides of
the clypellus straight (Fig. 12), and the
midtibial setal formula 4.3 or 4.4. A few speci-
mens examined (Stainback Highway) are
tinged with scarlet and have the abdomen and
thoracic stemites marked with dark brown.

Balclutha usitata Namba
Fig. 27

Balclutha usitata Namba. 19.56: 104-105. Figs. 2a-b.
Holot\"pe i . Kauai (BPBM) [examined].

Host. — Cyathodes (Gramineae).

Distribution. — Endemic (Kauai. Oahu,
Molokai. Maui. Hawaii .

Material E.\.\MiNED — Holot\-pe 6 , Kauai,
Kainamanu, vii.1952, 3,800' (BPBM). Nu-
merous specimens from throughout its range
BMNH. BPBM).

Renl\RKS. — This species can be distin-
guished by its small size (d , 2.6-2.9 mm, 9,
2.8-3.1 mm), its overall yellow or greenish
\ellow color without brown markings on the
dorsum of the abdomen, the 3.3 midtibial
setal formula, and the angularly cur\-ed aedea-
gal shaft (Fig. 27). Some specimens have
stramineous markings similar to but paler
than in volcanicola (Figs. 5, 7 .

Balclutha timberlakei (Osbom)
Figs. 13-15, 22

Nesosteles timberlakei Oshom. 1935: -59-60, Figs. 25a-d.

Holot>pe 6. Oahu BPBM [examined].
Balclutha timberlakei iOsbom(: Zimmerman 1948: 87,

Figs. 7e. 24c: Namba 19.56: 108-109, Figs. 9a-b.

Host. — Eragrostis variabilis (Gramineae).

Distribution — Endemic (Kauai. Oahu,

Great Basin Naturalist Memoirs

No. 12

Molokai, Lanai, Maui, Hawaii).

Material E>L\MINED. — Holotype S, Oahu
(BPBM). Oahu: \S (paratype) Palolo Val. on
Eragrostis variabilis (BMNH). Maui: Id,
Waikapu Val. (BPBM). Hawaii: M, 29, Po-
hakuloa, 1,800 m (BPBM).

Remarks. — This species can be distin-
guished by its small size ((5, 3.0 mm, 9 , 3.4
mm), elongate head that is distinctly wider
than the pronotum (Fig. 15), general yellow
color without brown markings on the dorsum
of the abdomen, 3.3 midtibial setal formula,
robust and slightly sinuate shaft of the aedea-
gus (Fig. 22), and its elongate connective,
which is longer than the style (Figs. 13, 14).

Balclutha phoxocephala Namba
Fig. 23

Balclutha phoxocephala Namba, 1956: 109, Fig. 10.
Holotype 6, Kauai (BPBM) [examined].

Host. — Cyathodes, Eragrostis variabilis
(Gramineae).

Distribution. — Endemic (Kauai, Oahu,
Molokai).

Material e.xamined. — Holotype 6 , Kauai:
Nualolo, viii.1925, on Cyathodes (BPBM).
Kauai, Oahu, Molokai (96, 59, paratvpes,
BPBM).

Remarks. — This species can be distin-
guished by its small size {6 , 2.4-2.6 mm, 9 ,
2.7-3.0 mm), 3.3 midtibial setal formula, and
robust aedeagus (Fig. 23). Some specimens
have stramineous markings similar to but
paler than in volcanicola.

A series of specimens from Hawaii, Pohaku-
loa, 6,300' (BPBM), may be this species but
have the aedeagal shaft more robust.

Balclutha incisa hospes (Kirkaldy), n. stat.

Figs. 28-40

N esosteles hebe \'dr . hospes, Kirkaldy. 1910: 574; Osborn
1935: 57, Figs. 24a-b. Lectotype 9, Hawaii
(BPBM), designated by Zimmerman (1948: 85)
[examined].

N esosteles hospes TimberVdke, 1918: .381.

Balclutha hospes (Kirkaldy): Zimmerman, 1948: 85, Fig.
24b, Namba 1956: 107, Figs. 6a-b.

Host. — Chloris radiata, Panicum piirpur-
ascens (Gramineae).

Distribution. — Cosmopolitan and through-
out the Hawaiian islands.

Material examined, — Lectotype 9 ofNe-
sosteles hebe var. hospes, Hawaii: Kauai
(BPBM). Numerous specimens from Kauai,
Oahu, Molokai, and Hawaii (BPBM, BMNH).

Remarks. — ^This subspecies can be distin-
guished by the head being slightly wider
than the pronotum, the short vertex (Fig. 28),
the ocelli being only 1.0-1.5 times their own
diameter from the corresponding eye, the
coronal suture visible in facial view (Fig. 29),
distinctively shaped aedeagus (Figs. 39, 40),
and the brown markings on the female pre-
genital sternite (Fig. 36). It is similar in exter-
nal appearance to B. saltuella (see Remarks
under salt uella).

This subspecies, which occurs in Hawaii,
the Marquesas Islands, and the Americas, dif-
fers from the nominate subspecies in the Old
World by the more laterally directed ventral
process of the aedeagus. This distinction sug-
gests that the species may have reached
Hawaii from the Americas rather than the
Pacific. Contrary to Timberlake's data (1918),
the shape of the pygofer process (Fig. 37) is
similar but variable in both subspecies.

Balclutha saltuella (Kirschbaum)
Figs. 41-46

Jassus (Tharnnotettix) saltuellus Kirschbaum, 1868; 86.
Holotype 9 , Germany (Museum Wiesbaden) [ex-
amined].

Balclutha beardsleyiNamhd, 1956: 110, 112, Figs, lla-b.
Holotype d, Hawaii (BPBM) [e.xamined], syn.
nov.

Host. — Unknown in Hawaii; elsewhere:
Agrostis (Gramineae) and Gossypium (Mal-
vaceae) (Knight 1987).

Distribution. — Cosmopolitan and on Oahu.

Material e.xamined. — Holotype 9 Jassus
{Tharnnotettix) saltuellus, Germany (Mu-
seum Wiesbaden). Holotvpe 6 B. beardsleyi,
Oahu: Honolulu, i. 1955 (BPBM). Oahu: 16,
39, Honolulu; 19, Ewa(BPBM).

Remarks. — This species is similar to B. in-
cisa in external appearance but differs from
this and other Hawaiian species by its reduced
number of foretibial spines (1.1), its medially
directed, lateral frontal sutures, and the
short, broad rostrum in facial view (Fig. 44). It
differs from incisa, in addition, by having the
prosternum dark brown (Fig. 43), the female
pregenital sternite with a medial, triangular
lobe on the posterior margin, and the disc
devoid of brown markings.

Nesolina Osborn

NesoHna Osborn, 1935: 60. Type species: Nesolina lineata
Osborn, bv original designation: Zimmerman

1948: 87.

1988

Knicht, Webb: Hawaiian Endemics in Balclutha

Figs. 28-46. Balclutha species. 28-40, B. incisa hospes: 28, head and thorax, dorsal view; 29, face; 30, apex of left
hind femur; 31, right forewing; 32, right hindwing; 33, apex of second valvulae, ventral view; 34, apex of left midtibia;
35, apex of left foretibia; 36, female pregenital sternite; 37, ventroposterior corner of male pygofer lobe, lateral view;
38, left second valvula, lateral view (toothed area between broken lines); 39, aedeagus, left lateral view; 40, aedeagus,
posterior view. 41-46, B. saltuella: 41, aedeagus, left lateral view; 42, connective, dorsal view; 43, head and thorax, left
lateral view; 44, face; 45, right style, dorsal view; 46, posteroventral corner of male pygofer lobe.

Remarks. — ^This endemic Hawaiian genus
can be distinguished by its long head and

distinctive markings, its large aedeagus with-
out processes, and the plumose macrosetae of

Great Basin Naturalist Memoirs

No. 12

Figs. 47-53. Balclutha lucida: 47, head and thorax, dorsal view; 48, left style, dorsal view; 49, male pygofer, left
lateral view; 50, left subgenital plate, ventral view; 51, aedeagus, left lateral view; 52, connective, dorsal view; 53,
distribution.

the legs and genitalia. The foretibia and
midtibia each have one long, and one short,
stout apical seta. Setal formulae: foretibia,
1.3; midtibia, 3.3; hind femur, 2.1.1. (see also
Remarks under "Relationship of Balclutha to
Other Hawaiian Macrostelini").

Nesolina lineata Osborn

Figs. 54-66

Nesolina lineata Osborn, 1935: 60-61, Figs. 26a-c: Zim-
merman 1948: 88, Figs. 7d, 8e-f, 24d. Lectotype
6 , Oahu (BPBM), here designated [examined].

1988

Knight, Webb: Hawaiian Endemics in Balclutha

Figs. 54-66. Nesolina lineata: 54, head and thorax, dorsal view; 55, face; 56, apex of left foretibia; 57, right forewing;
58, right hindwing; 59, immature, head and thorax, dorsal view; 60, apex of first valvulae; 61, aedeagus, left lateral view;
62^ aedeagus, posterior view; 63, left second valvula, lateral view (toothed area between broken lines); 64, male genital
segment, left lateral view; 65, apex of left style, lateral view; 66, subgenital plates, dorsal and ventral view, valve,
connective, and left style, dorsal view.

Host. — Eragrostis variabilis (Gramineae).
Distribution. — Endemic (Oahu, Hawaii).

Material examined. — Lectotype d, Oahu:
Diamond Head, 18. ii. 1917, W. M. Giffard.

100

Great Basin Naturalist Memoirs

No. 12

Oahu: IM, 89, 5 immatures, Diamond
Head, various dates (paralectotypes) (BPBM).
Hawaii: 16, Kau; 16, Pohakuloa, on bunch
grass (BMNH).

Balolina, n. gen.

Type species; Macrosteles kilatieae Kirkaldy.

Length. — 3.6-4.3 mm.

Yellow to greenish yellow, variably marked
with brown. Head slightly to distinctly nar-
rower than pronotum. Vertex triangular, me-
dial length approximately 1.5 times length
next to eye, foremargin broad to narrowly
parabolic. Ocelli marginal, visible dorsally,
situated approximately midway between eye
and midline. Face slightly wider than long,
lateral margin strongly sinuate below eye.
Clypeus narrow, lateral margins incurved at
level of eye. Transclypeal suture indistinct.
Clypellus narrow, slightly expanded to near
apex, extended well beyond margin of face.
Laterofrontal sutures extending one-half dis-
tance to corresponding ocellus. Pronotum 1.5
times length of vertex, lateral margins short.
Forewings elongate, appendix wide, extend-
ing around first and second apical cells; outer
subapical cell absent; inner subapical cell
open basally. Hindwing with veins r and m
fused distally. Forefemora with two stout api-
cal setae. Foretibia setal formula 1.3. Mid-
femora with one long, and one moderately
long, stout apical seta. Midtibia setal formula
usually 3.3. Hind femora setal formula usually
2.2.1. Base of hind tarsomere sulcate.

Male. — Pygofer narrowly rounded posteri-
orly; a group of submarginal macrosetae pos-
teriorly. Subgenital plates gradually tapered
to lightly sclerotized, fingerlike apex; a multi-
seriate row of macrosetae along ventrolateral
margin. Styles with apical process tapered to
acute apex in ventral view, apex footlike in
medial view; lateral lobe well developed; in-
ner basal apophyses short. Connective Y-
shaped; stem short, equal in length to arms;
articulating with aedeagus. Aedeagus with
shaft short and robust, cylindrical; a pair of
short processes apically; gonopore apical on
ventral surface; basal apodeme short.

Female. — First valvulae fused. Second
valvulae evenly tapered to acute apex; teeth
very fine, extending over distal half of dorsal
margin and onto apex of ventral margin.

Remarks. — This genus is similar in appear-

ance to Balchitha and Nesoclutha but can be
distinguished from these genera by the ab-
sence of hairy macrosetae, shorter laterofron-
tal sutures, indistinct transclypeal suture, the
ocelli situated more distant from the eyes, the
subgenital plates with multiseriate macrose-
tae, and the aedeagus with a pair of apical
processes. It differs from Balchitha also by the
other characters noted in the key.

Balohna kilaueae (Kirkaldy), n. comb.

Figs. 67-83

Macrosteles kilaueae Kirkaldy, 1910; 575. Holotype 9,

Hawaii (BPBM) [examined].
Balchitha kilaueae (Kirkaldy); Zimmerman 1948; 86,

Figs, 7a, Ud-f; Namba 1956; 103-104, Figs. la-b.

Host. — Cihotium chamissoi and C. men-
ziesii (Dicksoniaceae) (ferns).

Distribution. — Endemic (Kauai, Oahu,
Molokai, Maui, Hawaii).

Material examined. — Holotype 9,
Hawaii, Kilauea, vii.1906, RCL Perkins. Nu-
merous specimens from throughout its range
(BPBM, BMNH).

Remarks. — The specimens examined
varied in color and in the length of the vertex.
Brown markings are often present on the dor-
sal and ventral surfaces of the abdomen and
sometimes on the thorax, the clypeus, the
basal two-thirds of the ovipositor, the basal
half of the corium of the forewing, and the
hind legs. One specimen examined from
Maunawainui Valley, Molokai, has the clav-
us of the forewing pale scarlet. Setal formu-
lae: foretibia, 1.3; midtibia, usually 3.3 but
sometimes 4.3, 4.4, or 5.3; hind femur, usu-
ally 2.2.1, sometimes 2.1.1.1, 2.2.1.1, or
2.1.2.1.1.

This species is similar in external appear-
ance to Balchitha nambai with which it may
be confused.

Acknowledgments

We thank Keith Arakaki (BPBM) for the
loan of material under his care.

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1988

Knight. Webb: Hawaiian Endemics in Balclutha

101

Figs. 67-83. Balolina kilaueae: 67, head and thorax, dorsal view; 68, face; 69, apex of left femur; 70, right forewing;
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Zimmerman, E C 1948. Insects of Hawaii. Vol. 4. Ho-
moptera: Auchenorrhyncha. University of Hawaii
Press, Honolulu. 268 pp.

COLLADONUS AND RELATED GENERA OF MEXICO AND CENTRAL AMERICA
WITH NEW TAXA AND SYNONYMY (HOMOPTERA: CICADELLIDAE)

M. W. Nk'lson'

Abstract. — A study of the genus CoUadonus and related genera of Mexico and C^entral Ameriea was based almost
exclusively upon the examination of nearly the entire type series of all the Idiudomis species described by the late Dr.
D. M. DeLong. All species referable to CoUadonus are redescribed, and illustrations of the male genitalia are given for
the first time. A key to the species of Mexico and Central America is also included. A brief discussion of the distribution
and phylogeny of the group is given.

Among 35 species described by DeLong in Idiodonus , 21 belong in CoUadonus, 1 is assigned to OUarianus Ball, 1 to
Paratanus Young, 2 to Bonncyana Oman, 2 are retained in Idiodonus , 5 are relegated to four proposed new genera, and
the remaining 3 have uncertain generic position. One new combination in CoUadonus is proposed, and 16 names are
treated as new synonyms. Eight new combinations are proposed in the treatment of related genera. New genera
include: ParacoUadonus, Paracrassana, Paranurenus, and Jaacunga. The subgenus Angulanus DeLong is elevated to
generic rank.

The genus CoUadonus Ball was first revised
by Nielson (1957), who treated primarily the
Nearctic species north of Mexico and "one
known single Palearctic species. After the ma-
jor part of the revisionary work was finished, a
number of Mexican leafhoppers described by
DeLong (1946) in the genus Idiodonus Ball
were studied, and several species were found
referable to CoUadonus . Twelve species were
listed as new combinations and treated as in-
certae sedis in the appendix of my 1957 paper.
Two species-level names were suppressed as
new synonyms of an older name. Time did not
permit a more thorough study of suspected
additional synonymy and possible dissociation
of the sexes between the female holotype and
male allotype of several species in this group.

The descriptions of nearly all of De Long's
Idiodonus species were based on females, and
a female specimen was designated as the holo-
type in nearly all cases. Illustrations were lim-
ited to a dorsal view of the head and thorax,
and a ventral view of the female seventh ster-
num, which with color patterns formed the
basis of his classification. In a much later pa-
per, DeLong (1983) described additional spe-
cies of Idiodonus from Mexico and Bolivia;
then he (DeLong 1984) prepared a revised
key to all the known species in the New
World, separating them on the basis of color
patterns and configuration of the female sev-
enth sternum.

Although the female seventh sternum alone
is a useful character for separating some spe-
cies o( CoUadonus, its utility is enhanced only
in combination with male genital characters.
This is particularly true in certain species that
show similarities in male characters that re-
quire support by associated female seventh
sternal characters to differentiate these spe-
cies.

In this study the entire type series, with the
exception of a few specimens, of all species of
Idiodonus described by DeLong were exam-
ined with attention given to association of the
female holotype and her counterjDart male al-
lotype or male paratype specimen. A few
cases of improper sex association were found,
and these are discussed below. All species
found referable to CoUadonus are re-
described, and illustrations of the male geni-
talia are given for the first time. A key to the
species of Mexico and Central America is also
included. Six new species of CoUadonus are
described from that region. One new combi-
nation is proposed, those treated as new com-
binations in my earlier paper are confirmed,
and 16 names are treated as new synonyms
under their respective nominate species-level
name. CoUadonus delongi described by Lin-
navuori (1959) from Panama and based on an
apparent mislabeled and misdetermined fe-
male specimen was treated as a new synonym
of CoUadonus montanus (Van Duzee) by Niel-
son (1966).

'Monte L. Bean Life Science Museum, Brigham Young University, Provo, Utah 84602.

103

104

Great Basin Naturalist Memoirs

No. 12

Among 35 species described in Idiodonus
by DeLong, 21 belong in CoUadonus (includ-
ing 14 species treated in my earlier work), 1 is
assigned to Ollarianus Ball, 1 to Paratamis
Young, 2 to Bonneyana Oman, 2 are retained
in Idiodonus, 5 are relegated to four proposed
new genera, and the remaining 3 have uncer-
tain generic position. One species, /. plum-
meri DeLong, is considered the only valid and
correctly placed species among all those de-
scribed in Idiodonus by DeLong. Idiodonus
bakeri DeLong is suppressed as a new syn-
onym of plummeri. DeLong treated 6 older
species of Idiodonus in his 1946 paper, 5 of
which were subsequently assigned to three
other genera by Oman (1949). Descriptions of
four new genera, ParacoUadonus, Paracras-
sana, Paranurenus, and Jaacunga, are pro-
vided with redescriptions and illustrations of
the species assigned to their respective genus.
The subgenus Angulanus DeLong is elevated
to generic rank.

Distribution of Colladonus . — The origin
of the genus appears to have been centered in
the southern highland range of the Nearctic
region (Rocky Mountain subregion) in Mexico
where radiation northward produced the bulk
of species in United States. Only one species
is known to occur as far south as Panama, and
one is widespread in the Palearctic region.
Fifteen species occur in Mexico proper, only
one of which ranges into Arizona. Among 44
species in United States (including Alaska)
and Canada, seven occur marginally in Mex-
ico (Bliven 1954, Nielson 1962, 1966). Most of
the species of CoUadonus occur on trees and
shrubs in the mountainous regions of western
United States and Canada.

Genera closely allied to CoUadonus {Nigri-
donus Oman, Caladonus Oman, Bonneyana
Oman, and two new genera) are known only
from Mexico and the extreme southwestern
mountain areas of the United States, lending
support to the Mexican origin of the genus
CoUadonus.

With respect to the New World distribution
of Idiodonus, it is doubtful that the genus
occurs south of Mexico. Members of the
genus are more common in the United States
than in Mexico.

Considerations of phyloceny— Nearly
all female CoUadonus of species that occur
north of Mexico have a spatulate process on
the caudal margin of the seventh sternum.

None of the species in Mexico and Central
America possess this structure, suggesting
that it is a derived (apomorphic) character.
The spatulate process thus far has attained its
highest development in C. furculatus (Os-
born) that occurs in the eastern United States
and Canada. There does not appear to be any
correlation between the presence of this
structure and male genital structures, indicat-
ing that its development was independent of
the evolutionary development in male struc-
tures. However, there appears to be a correla-
tion between the absence of the spatulate pro-
cess on the female seventh sternum and a
short, male pygofer spine among the species
(United States, Canada, and Palearctic re-
gion) that possess these characters, viz., beUi,
youngi, hrunneus, and torneeUus. This rela-
tionship does not hold true for species in Mex-
ico and Central America, suggesting that the
northern population evolved separately from
its sister population in the United States and
Canada that does not exhibit this relationship.
About half of the CoUadonus species have
crowns that are produced, which is consid-
ered a more primitive condition. Associated
with this character is the relative position of
the gonopore on the shaft of the aedeagus. A
gonopore basad of the midlength of the aedea-
gal shaft is considered more primitive, where-
as a position distad of middle is more ad-
vanced. About 82% of the United States and
Canadian species bearing a basal gonopore
have a produced crown, whereas all of those
species bearing a distal gonopore have a short,
rounded crown. Among the Mexican and
Central American species, nearly all species
with a basal gonopore have a produced crown,
whereas all species with a distal gonopore
have a short, rounded crown. Each of those
sister groups apparently evolved indepen-
dently.

Key to Males oi Colladonus of Mexico
and Central America

1. Pronotum with yellow or ivory, (sometimes

pale) transverse band 2

— Pronotum without such band 13

2(1). Pygofer with a ver\- short, exposed spine on
caudal margin, exposed length about twice as
great as exposed basal width 3

— Pygofer with moderately long to very long,
exposed spine on caudal margin, exposed
length nearly three times to more than five
times as great as exposed basal width 8

1988

NiELSON; COLLADONUS OF MEXICO AND CENTRAL AMERICA

105

14(13).

15(14).

3(2). Pygofer spine near middle of caudal margin;
connective short, not reaching to apex of

style; aedeagus with bifurcated processes

long, reaching to midlength of shaft 4

— Pygofer spine distinctly hasad of middle of
caudal margin; connective long, reaching to
apex of style; aedeagus with bifurcated pro-
cesses short, not reaching to midlength of

shaft albocinctus (DeLong) 13(1).

4(3). Pygofer spine near middle or just below mid-
dle of caudal margin 5

— Pygofer spine distinctly above middle of can-
dal margin belli (Uhler)

5(4). Pygofer with caudoventral margin smooth;

style with apical stylar spine 6

— Pygofer with caudoventral margin toothed or
serrate; style with subapical stylar spine ....
serratus, n. sp.

6(5). Aedeagus with bifurcated processes long, ex-
tending beyond gonopore in lateral view .... 7

— Aedeagus with bifurcated processes short, not

reaching to gonopore in lateral view

fasciaticollis (Stal)

7(6). Style with apical shaft broad and lateral mar-
gins parallel, stylar spine apical and project-
ing laterad ultimus, n. sp.

— Style with apical shaft narrowed distally, lat-
eral margins not parallel, stylar spine apical
and projecting laterodistad . . . tolucensis, n. sp.

8(2). Pygofer spine with exposed length not more

than 4 times exposed basal width 9

— Pygofer spine with exposed length more than
5 times exposed basal width 11

9(8). Connective short, not reaching to apex of
style; aedeagus in lateral view with bifurcated
process short, reaching to about midlength of
shaft 10

— Connective long, reaching to about apex of
style; aedeagus in lateral view with bifurcated
processes long, extending beyond midlength
of shaft dampfi (DeLong)

10(9). Pronotum with narrow, black band on ante-
rior margin and narrow, black, transverse
band on middle; style with lateral margins of
distal shaft smooth bicinctus (DeLong)

— Pronotum with narrow, black band on ante-
rior margin but without medial, black, trans-
verse band; style with inner lateral margin of
distal shaft toothed claustriis (DeLong)

11(8). Pygofer with caudoventral margin and
pygofer spine smooth, without serrations or
teeth 12

— Pygofer with caudoventral margin serrate or

toothed, pygofer spine toothed basally

verecundus (DeLong)

12(11). Pygofer with spine arising about middle of
nearly truncate caudal margin; connective
very long, extending beyond apex of style;
aedeagus with bifurcated processes short,

16(14)

nearly reaching to midlength of shaft

titulus (DeLong)

Pygofer with spine arising from apex of dis-
tally produced caudoventral margin; connec-
tive short, extending nearly to apex of style;
aedeagus with bifurcated processes long, ex-
tending beyond midlength ol shaft

sirifiularius, n. sp.

Pronotum never with yellow, longitudinal
band; claval veins not deeply marked with
yellow, if yellow, very pale or ivory 14

Pronotum with yellow, lontiituclinal band,
claval veins deeply marked with yellow ....
trabilis, n. sp.

Pygofer spine very long, exposed length more
than 5 times exposed basal width 15

Pygofer spine short, exposed length not more

than 3 times exposed basal width 16

Crown with two handlebar-shaped markings
on anterior margin; aedeagus with very long,
reflexed, bifurcated processes in lateral view,

processes parallel in dorsal view

clathrus (DeLong)

Crown with as many as 8 black spots on ante-
rior margin, some spots sometimes con-
nected, if so, never handlebar shaped; aedea-
gus with short bifurcated processes nearly
reaching midlength of shaft in lateral view,

processes crossing over in dorsal view

beameri (Ball)

Pygofer with caudal margin smooth 17

Pygofer with caudal margin serrate or toothed

17(16).

18(16).

Pygofer with very short spine arising from
caudoventral margin; aedeagus with gono-
pore at about middle of shaft; connective long,

reaching to about apex of style

albocinctus (DeLong)

Pygofer with moderately long spine arising
from middle of caudal margin; aedeagus with
gonopore basad of middle shaft; connective

short, not reaching apex of style

tessellatus. n. sp.

Pygofer with caudoventral margin produced
distally, small, robust spine arising from cau-
dodorsal angle, projecting dorsally; style with
distal shaft long and narrow, serrate on inner
lateral margin; aedeagus with gonopore at

middle of shaft. . anademus (DeLong)

Pygofer with obtusely angled caudal margin,

very small spine arising from apex of angle and
projecting distad; style with distal shaft robust
and short, smooth on inner lateral margin;
aedeagus with gonopore basad of middle of
shaft incidus (DeLong)

Colladonus albocinctus (DeLong)
Figs. 1-5

Idiodonus albocinctus DeLong 1946;22 [Holotype 9

(OSU) (examined).]
Colladonus albocinctus: Nielson 1957:51

106

Great Basin Naturalist Memoirs

No. 12

Figs. 1-5. Colladonus albocinctiis (DeLong): 1, male pygofer, lateral view; 2, connective and right style, dorsal view;
3, aedeagus, lateral view; 4, apex of right style, enlarged dorsal view; 5, apex of aedeagus and distal processes, enlarged
dorsal view.

Idiodonus albocinctus: Metcalf 1967:1286

Idiodonus albocinctus: DeLong 1984:10

Idiodonus nigridens DeLong 1946:29 [Holotype 9 (OSU)

(examined).] New synonymy

Colladonus nigridens: Nielson 1957:51

Idiodonus nigridens: Metcalf 1967:1301

Idiodonus nigridens: DeLong 1984:10

Idiodonus sexpunctatus DeLong 1983:90 [Holotype 9

(OSU) (examined).] New synonymy

Idiodonus sexpunctatus: DeLong 1984:10

Length. — IVIale, 4.9-5.0 mm; female, 5.0-
5.3 mm.

Color variable from absence to presence of
narrow, ivory, transverse band about middle
of pronotum, with two black spots on anterior
margin of crown and absence or presence of

narrow, black, transverse band on disc of
crown. Similar in general habitus to verecun-
dus (DeLong) and in male genitalic characters
to fasciaticoUis (Stal).

Head with anterior margin produced, apex
obtusely angulate.

Male. — Pygofer in lateral view with very
short, exposed caudoventral spine arising be-
low middle of caudal margin, basal nine-
tenths of spine sclerotized and fused to
pygofer wall (Fig. 1); aedeagus in lateral view
with bifurcated distal processes less than half
as long as aedeagal shiift, not reaching
gonopore (Fig. 3), crossing over in dorsal view
(Fig. 5), gonopore at about midlength of shaft;

1988

NiELSON: COLLADONUS OF MEXICO AND CENTRAL AMERICA

107

connective long, extending to about apex of
style (Fig. 2); style with distal shaft broad,
lateral margins smooth, stylar spine broad
and bluntly pointed, projecting laterodistallv
(Fig. 4).

Female. — Seventh sternum produced dis-
tally to form broad medial lobe, with median,
shallow notch on its caudal margin, configura-
tion of lobe variable, sometimes with small
lobe at base of notch.

Distribution. — This species has been col-
lected in the Mexican states of Distrito Fed-
eral, Michoacan, Morelos, and Hidalgo.

Host. — Some specimens were collected in
pine forests at elevations of 7,500 feet from
July to October (DeLong 1946).

Remarks. — The configuration of the female
seventh sterna of the holotypes of nigridens
and sexpunctatiis is similar, with some intra-
and interpopulation variation in the angle of
the posteriolateral margins. The abdomen of
the female holotype oi albocinctus is missing,
but comparisons were made with paratype
female specimens after they were associated
with holotype and with DeLong's illustration
of the seventh sternum. Although the male
genital structures of the allotypes of albocinc-
tus and nigridens are nearly identical, the
latter is more deeply sclerotized than the for-
mer. The sexes of both species appear to be
properly associated.

The female holotype of sexpunctatus is
identical in general habitus with the female
holotype oi nigridens, but both species differ
markedly in color patterns from albocinctus.
The latter species has a distinctive pronotal
band that is absent in the specimens of the two
former species. The male genital structures,
however, are identical among these popula-
tions, which suggests that two infraspecific
entities may be involved. Such action is de-
ferred until more material is collected and
studied.

Colladonus albocinctus is distinguished
homfasciaticollis by the position of the pygo-
fer spine, which is distinctly basad of the mid-
dle of the caudal margin of the pygofer.

Colladonus belli (Uhler)
Figs. 6-10

Jassus belli Uhler 1877:471 [Holotype 9 (USNM) (exam-
ined).]
Thamnotettix belli: Van Duzee 1892:306
Colladonus belli: Ball 1936:58
Idiodonus belli: Medler 1943:18

Colladonus belli: Nielson 1957:38
Colladonus belli: Metcalf 1967: 1245
Thamnotettix semipullatus Van Duzee 1892:306
Colladonus semipullatus: Oman 1949:125
Thamnotettix fiilletti Van Duzee 1892:306
Colladonus gilk'tti: Oman 1949:125
Thamnotettix sonorae (;illette & Baker 1895:100
Colladonus sonorae: Oman 1949:125

Length. — Male, 4.0-4.4 mm; female, 4.4-
5.2 mm.

A well-marked species from pale yellow to
deep yellow with two black spots on anterior
margin of crown and sometimes with a black,
transverse band on disc of crown; pronotum
with pale yellow to deep yellow, transverse
band; forewings with veins pale to yellow;
color intensity highly variable. Similar to fas-
ciaticollis in general habitus but with distinc-
tive male genitalia.

Head with anterior margin not produced,
apex rounded.

Male. — Pygofer in lateral view with short,
stout spine near or above middle of caudal
margin, spine projecting dorsally (Fig. 6);
aedeagus in lateral view with long, bifurcated
distal processes extending beyond midlength
of shaft (Fig. 8), crossing over in dorsal view
(Fig. 10), gonopore about middle of shaft; con-
nective short, not reaching apex of style (Fig.
7); style with moderately broad stylar shaft,
inner lateral margin serrate, outer one
smooth, stylar spine moderately long, pro-
jecting laterally (Fig. 9).

Female. — Seventh sternum with truncate
caudal margin, shallow, U-shaped excision
medially.

Distribution. — This species is widespread
in mountainous areas from Mexico north to
British Columbia. In Mexico the author col-
lected specimens in the state of Chihuahua. It
has been reported in seven western states in
the United States and in the province of
British Columbia (Nielson 1957).

Host. — Adults have been swept from a va-
riety of trees and shrubs by the author. Sev-
eral specimens were taken from Monarda sp.
near citriodora Cerv. in Chihuahua state in
Mexico by the author.

Remarks. — Colladonus belli may repre-
sent a species complex that occupies a rather
narrow, longitudinal range in western North
America. Specimens north of Utah and
Colorado are almost uniformly yellow-green
without a dark band across the crown,
whereas southern populations are more

108

Great Basin Naturalist Memoirs

No. 12

Figs. 6-10. CoUadomts belli (Uhler): 6, male pygofer, lateral view; 7, connective and right style, dorsal view; 8,
aedeagus, lateral view; 9, apex of right style, enlarged dorsal view; 10, apex of aedeagus and distal processes, dorsal
view.

deeply marked with yellow and black mark-
ings. The position of the pygofer spine, al-
though similar in configuration, varies from
distad of the middle of the caudal margin in
northern populations to the middle of the cau-
dal margin in southern populations. No varia-
tion in the configuration of the female seventh
sternum was evident, however.

Much additional collecting over the entire
range of this species complex is needed. Pop-
ulations are not commonly collected, and
when found, specimens are usually few in
number. A reexamination of the type speci-
mens of belli, semipullatus, ^illettei, and
sonorae is recommended for any future study.

From fasciaticollis, to which it is similar,
belli can be distinguished by the short pygofer
species has been made, nor has any type of

die to upper middle portion of the caudal
margin.

Colladonus serratiis, n. sp.
Figs. 11-14

Length. — Male, 4.8-5.0 mm; female, 5.4
mm.

Color dark brown to blackish; crown yellow
with two large, (juadrate, black spots on ante-
rior margin, disc with broad, black, transverse
band; pronotum with yellow, narrow to
broad, transverse band; forewings black ex-
cept for transculent area along middle of
costa, veins yellow; face yellow with black
markings.

Head with anterior margin produced, apex
obtusely angled.

Male. — Pygofer in lateral view with
broadly roimded caudal margin, caudoventral

1988

NiELSON: COLLADONUS OF MEXICO AND CENTRAL AMERICA

109

Figs. 11-14. CoUadonus serratus, n. sp.: 11, male pygofer, lateral view; 12, connective, right style, and aedeagus,
dorsal view; 13, aedeagus, lateral view; 14, apex of right style, enlarged dorsal view.

margin toothed or serrate, short spine above
middle of caudal margin, directed dorsad
(Fig. 11); aedeagus in lateral view narrow
along major portion of shaft with moderately
long, distal, bifurcated processes reaching to
about middle of shaft (Fig. 13), processes
broad medially, gradually tapered distally,
crossing over in dorsal view (Fig. 12), gono-
pore basad of middle of shaft; connective
short, not reaching apex of style (Fig. 12);
style with broad stylar shaft, stylar spine sub-
apical, small, sharply pointed (Fig. 14).

Female. — Seventh sternum with truncate
caudal margin, small tooth medially.

HoLOTYPE (male). — Mexico: Mex., 7 miles
east of Amecameca, 9,300 ft, 31. VI. 1974, C.
W. and L. O'Brien and Marshal (USNM).
Allotype female, same locality as holotype,
10,000 ft, 17. VIII. 1982, C. W. and L. O'Brien
and G. Wibner (USNM). Paratypes: one
male, same data as female allotype (in au-
thor's collection); one male, Mexico, #16, 7
miles southeast of Amecameca, 9,000 ft,
17. VIII. 1969, George W. Byers (UK).

Remarks. — ^This is a rather unique species
that is not related to other species oi CoUa-

donus in Mexico. The combination of the sub-
apical stylar spine and toothed caudovcntral
margin of the pygofer will readily distinguish
it from all other known species in the region.

CoUadonus fasciaticollis (Stal)
Figs. 15-22

J assiis fasciaticollis Stal 1864:86 [Holotype 9 (NM) (ex-
amined).]

Thamnotettix fasciaticollis: DeLong 1946:13

CoUadonus fasciaticollis: Nielson 1957:37

Thamnotettix fasciaticollis: Metcalf 1967:756

Idiodonus tubtdus DeLong 1946:22 [Holotype 9 (OSU)
(examined).]

CoUadonus tubulus: Nielson 1957:37

Idiodonus tubulus: Metcalf 1967:1303

Idiodonus tubulus: DeLong 1984:10

Idiodonus diserus DeLong 1946:24 [Holotype 9 (OSU)
(examined).]

CoUadonus diserus: Nielson 1957:37

Idiodonus diserus: Metcalf 1967:1297

Idiodonus diserus: DeLong 1984:11

Idiodonus pravus DeLong 1946:24 [Holotype 9 (OSU)
(examined).] New synonymy

Idiodonus pravus: Metcalf 1967:1301

Idiodonus pravus: DeLong 1984:11

Idiodonus dampfi DeLong 1946:20 [In part, allotype 6
and 8 paratypes (OSU)]

Idiodonus dampfi: Linnavuori 1959:280 [Misdetermined
specimens.]

Great Basin Naturalist Memoirs

No. 12

Figs. 15-22. Colladonus fasciaticollis (Stal): 15, male pygofer of specimen from Costa Rica, lateral view; 16, male
pygofer of allotype oi tubulus , lateral view; 17, male pygofer of allotype oi disertis, lateral view; 18, male pygofer of
allotype of dampfi (= fasciaticollis), lateral view; 19, connective and right style, dorsal view; 20, aedeagus, lateral view;
21, apex of right style, enlarged dorsal view; 22, apex of aedeagus and distal processes, enlarged dorsal view.

Length. — Male, 4.4-4.6 mm; female,
4.8-5.0 mm.

A well-marked black and yellow species
usually, but sometimes paler in some speci-
mens. Similar in general habitus to belli and to
albocinctus in male genitalic characters but
distinct from both species.

Head with anterior margin not produced,
apex nearly rounded.

Male. — Pygofer in lateral view with short,
bluntly pointed spine arising below middle of
caudal margin (some variation in origin of
spine on caudoventral margin), basal 3/4 of
spine sclerotized and fused to pygofer wall
(Figs. 15, 16, 17, 18); aedeagus in lateral view
with moderately long and narrow (sometimes
broad), distal, bifurcated processes extending
to about middle of shaft (Fig. 20), crossing
over in dorsal view (Fig. 22), gonopore at

about middle of shaft; connective short, apex
not reaching to apex of style (Fig. 19); style
with tapered stylar shaft, lateral margins
smooth, stylar spine slanted laterodistally
(Fig. 22).

Female. — Seventh sternum with caudal
margin obtusely angled with shallow, U-
shaped, median emargination.

Distribution. — Colladonus fasciaticollis is
widely distributed from southern California to
Costa Rica. Its northern range into the United
States is restricted to the southern border of
California. It is common in southern Mexico
in the states of Distrito Federal, Veracruz,
Michoacan, and Puebla.

Host. — This species presumably occurs on
unspecified trees or shrubs in Mexico from
September to November at elevations from

1988

NiELSON:CoLLADONus OF Mexico AND Central America

111

5,000 to 10,000 feet (DeLong 1946). In Costa
Rica it was taken on Lippia herlandieri in
February (Nielson 1957).

Remarks. — In my earlier work (Nielson
1957), tubulus and diserus were transferred to
Colladonus and suppressed as synonyms of
fasciaticollis . Reexamination of the female
holotypes and comparison of them with au-
thentically determined female specimens of
fasciaticollis confirm that action.

The female holotype of pravus is identical
to its counteipart holotype specimens of the
above species. The male allotype of dampfi
from Bella Vista, Mexico (MB 366), which
apparently had been misidentified and im-
properly associated with the female holotype,
was associated with the male holotype of
pravus from Zitacuaro, Michoacan. Two of
five male paratype specimens and three fe-
male paratype specimens of dampfi from
Orizaba, Veracruz, were also associated with
pravus.

From albocinctus and belli, to which it has
similar genitalia, fasciaticollis can be distin-
guished by the pygofer spine that arises from
near the middle of the caudal margin, the
shorter aedeagus, and the stouter, bifurcated
distal processes.

Colladonus idtimus, n. sp.

Figs. 23-26

Length. — Male, 3.9-4.2 mm; female,
4.7-4.9 mm.

General color dark brown to black; crown
yellow with two black spots on anterior mar-
gin and narrow, black, transverse band be-
tween eyes; pronotum black with narrow, yel-
low, transverse band at middle; forewings
black with yellow veins, costal area hyaline;
face yellow with black markings. Similar in
general habitus to dampfi (DeLong) but with
distinctive male genitalia.

Head with anterior margin produced, apex
rounded.

Male. — Pygofer in lateral view with caudal
margin broadly rounded, pygofer spine short,
slightly longer than basal width, arising just
below midline of pygofer and projecting cau-
dodorsally (Fig. 23); aedeagus in lateral view
with narrow shaft, distal, bifurcated processes
long, more than 1/2 length of shaft and extend-
ing beyond gonopore (Fig. 25), processes of
uniform width except for tapered apex,
gonopore at about midlength of shaft; connec-

tive short, apex not reaching apex of style
(Fig. 24); style with broad stylar shaft, nearly
straight, stylar spine short, sharply pointed
distally and projecting laterally (Fig. 26).

Female. — Seventh sternum with caudal
margin nearly truncate, with small indenta-
tion medially.

Holotype (male). — Panama: Chiriqui,
Bambito, 1,400 m, 10. VI. 1976, Wolda and
Estribi(USNM).

Remarks. — Numerous specimens of this
species that were collected in Costa Rica and
preserved in alcohol were too badly distorted
and discolored to be used in the type series.
They were all taken at Cerro de la Muerte,
10,000 ft, I. VIII. 1966, S. L. Wood, and are in
the M. L. Bean Museum collection. A single
female specimen of this species from Yepo-
capa, Guatemala, XII. 1948, H. T. Dalmat,
was also examined.

This species thus far represents the south-
ernmost extremity of the range ofColladonus.
It can be distinguished from dampfi by the
shorter pygofer spine, by the gonopore,
which lies midlength of the aedeagal shaft,
and by the shorter connective.

Colladonus tolucensis, n. sp.
Figs. 27-30

Length. — Male, 4.5-4.6 mm; female, 5.0-
5.2 mm.

Color tan with black markings; crown with
two spots on anterior margin, black, trans-
verse band on middle of crown; pronotum
with broad, pale ivory, transverse band,
sometimes with broken black stripe on either
side of middle; forewings with pale veins,
clavus sometimes dark brown, and with dark
brown, longitudinal stripe above costal cell;
face deep tan with black markings on clypeus.
Similar in general habitus to several species
possessing broad pronotal band and in male
genital characters to idtimus.

Head with anterior margin produced, apex
obtusely angled.

Male. — Pygofer in lateral view with broad,
obtusely rounded caudal margin, very short
spine arising from middle of margin and some-
times hidden from lateral view, projecting
distally (Fig. 27); aedeagus in lateral view with
narrow shaft, distal, bifurcated processes
long, extending slightly beyond midlength of
shaft (Fig. 29), processes broad medially,
pointed distally, gonopore at about midlength

112

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No. 12

Figs. 23-26. CoUadonus ultimus, n. sp. : 23, male pygofer, lateral view; 24, connective, right style, and aedeagus,
dorsal view; 25, aedeagus, lateral view; 26, apex of right style, enlarged dorsal view.

of shaft; connective short, not reaching apex of
style (Fig. 28); style with tapered stylar shaft,
stylar spine small, directed laterodistallv (Fig.
30).

Female. — Seventh sternum with caudal
margin nearly truncate, with broad median
excavation.

HoLOTYPE (male). — Mexico: Distrito Fed-
eral, Toluca road, 26. IX. 1945, DeLong,
Hershberger, and Elliot (OSU), allotype fe-
male, Mexico City, D.F., 13.IX.1939, D. M.
DeLong (OSU). Paratypes: one male, same
data as holotype (in author's collection); one
female, same data as allotype (in author's col-
lection); 29 females, same data as allotvpe
(OSU).

Remarks. — From ultimus, to which it is
similar, tolucensis can be distinguished by the
unexposed pygofer spine, by the tapered sty-
lar shaft, and by the color patterns on the
pronotum.

CoUadonus dampfi (DeLong)

Figs. 31-34

kliodoniis dampfi DeLong 1946:20 [Holotype 9 (OSU)

(examined).]
CoUadonus dampfi: Nielson 1957:51
Idiodonus dampfi: Metcalf 1967:1297
Idiodomis dampfi: DeLong 1984:11

Length. — Male, 3.9-4.0 mm; female, 4.5
mm.

A well-marked, black and yellow species
with two distinct, black spots on anterior mar-
gin of crown, narrow, transverse, black band
on disc; pronotum with broad, black band
along anterior margin, broad, yellow, trans-
verse band medially, narrow, black band on
posterior margin; forewings with pale yellow
to deep yellow veins, cells brown to black.
Similar to other pronotal banded species and
to bicinctus (DeLong) in male genital charac-
ters.

Head with anterior margin not produced,
apex obtusely rounded.

Male. — Pygofer in lateral view with mod-
erately long, narrow spine arising from mid-
dle of caudal margin and projecting dorsally

1988

NiELSON: COLLADONUS OF MEXICO AND CkNTHAI. AMERICA

113

Figs. 27-30. Colladonus tohicensis, n. sp. : 27, male pygofer, lateral view; 28, connective, right style, and aedeagus,
dorsal view; 29, aedeagus, lateral view; 30, apex of right style, enlarged dorsal view.

(Fig. 31); aedeagus with distal, bifurcated pro-
cesses long, extending beyond midlength of
shaft (Fig. 33), crossing over in dorsal view,
gonopore basad of middle of shaft; connective
long, extending to about apex of style (Fig.
32); style with narrow stylar shaft, inner lat-
eral margin roughly dentate, stylar spine long
and projecting laterally (Fig. 34).

Female. — Seventh sternum with caudal
margin narrowed distally, shallow, U-shaped
indentation medially.

Distribution. — Colladonus dampfi is known
only from the Mexican states of Veracruz and
Morelos.

Host. — Recorded from unspecified trees
and shrubs from 3,000 to 9,000 feet during
September to November (DeLong 1946).

Remarks. — The type series of dampfi is
mixed. The female holotype and male allotype
were not properly associated; the latter is
identical to pravus, which is treated as a ju-
nior synonym oi fasciaticollis in this paper. A
male paratype oi dampfi from Cordoba, Vera-
cruz (type locality), was associated with the
female holotype of dampfi and used in illus-

trating the male genitalia. Five additional
male paratypes, three from Cuernavaca,
Morelos, and two from unspecified localities
(MB 95, Mexico, and MF 8523, Mexico) were
identical to the male paratype specimen asso-
ciated with the female holotype. The remain-
ing specimens in the type series are discussed
under fasciaticollis.

Colladonus darnpfi can be distinguished
from bicinctus by the short and more rounded
anterior margin of the head, by the longer
pygofer spine, and by the longer, distal, bifur-
cated aedeagal processes.

Colladonus bicinctus (DeLong)
Figs. 35-39

I diodomis bicinctus DeLong 1946:18 [Uo\otype 9 (OSU)

(examined).]
Colladonus bicinctus: Nielson 1957:51
Idiodonus bicinctus: Metcalf 1967:1287
Idiodonus bicinctus: DeLong 1984:10

Length. — Male, 4.2 mm; female, 4.5-4.8
mm.

A well-marked species of deep yellow with
black markings; crown with two triangular,

114

Great Basin Naturalist Memoirs

No. 12

Figs. 31-34. CoUadonus dampfi (DeLong): 31, male pygofer, lateral view; 32, connective, right style, and aedeagus,
dorsal view; 33, aedeagus, lateral view; 34, apex of right style, enlarged dorsal view.

black spots on anterior margin and narrow,
black, transverse band on middle of disc;
pronotum with narrow, black band along
anterior margin and narrow, black, transverse
band on middle transecting broad, yellow,
transverse band; forewings with veins yellow,
cells black. Similar in general habitus and
male genitalic characters to claustrus (De-
Long).

Head with anterior margin produced, apex
angled.

Male. — Pygofer in lateral view with mod-
erately long spine on middle of caudal margin,
spine directed caudodorsally (Fig. 35); aedea-
gus in lateral view with stout, distal, bifur-
cated processes reaching to about midlength
of shaft (Fig. 37), not crossing over in dorsal
view (Fig. 39), gonopore slightly basad of mid-
length of shaft; connective short, not reaching
apex of style (Fig. 36); style with narrow stylar
shaft, lateral margins smooth, stylar spine
very short and bluntly pointed (Fig. 38).

Female — Seventh sternum with caudal
margin shallowly and broadly concave, small
indentation medially.

Distribution — This species is known only
from the Mexican state of Distrito Federal.

Host. — According to DeLong (1946) the
species was collected at 7,500 feet from un-
specified shrubs or trees during a June-
September period.

Remarks, — ^This species was described
from the female holotype and a female para-
type specimen. However, a male paratype
specimen (MB 352, Mexico, A. Dampf, coll.),
which was without a name label and not cited
in the original description, was found among
the type series. The specimen was associated
with the female holotype and used in illustrat-
ing the genitalia.

The species can be separated from claus-
trus by the presence of a narrow, transverse,
black band on the middle of the pronotum, by
the narrower pygofer spine, by the shorter

1988

NiELSON: COLLADONUS OF MEXICO AND CENTRAL AMERICA

115

Figs. 35-39. Colladonus bicinctus (DeLong); 35, male pygofer, lateral view; 36, connective and right style, dorsal
view; 37, aedeagus, lateral view; 38, ape,\ of right style, enlarged dorsal view; 39, apex of aedeagus and distal processes,
enlarged dorsal view.

connective, and by the smooth-margined sty-
lar shaft.

Colladonus claustrus (DeLong)
Figs. 40-44

Idiodonus claustrus DeLong 1946:18 [Holotype 9 (OSU)

(examined).]
Colladonus claustrus: Nielson 1957:51
Idiodonus claustrus: Metcalf 1967:1289
Idiodonus claustrus: DeLong 1984:11

Length. — Male, 4.4 mm; female, 4.8-5.0
mm.

A well-marked species with color patterns
nearly identical to bicinctus except paler and
without median, transverse, black band on
pronotum. Similar in male genital characters
to bicinctus.

Head with anterior margin produced, apex
angled.

Male. — Pygofer in lateral aspect with mod-
erately long, stout spine on middle of caudal
margin, spine directed caudodorsally (Fig.
40): aedeagus with slender, bifurcated, distal
processes reaching to about middle of shaft,
apices not reaching gonopore (Fig. 42), cross-
ing over in dorsal view (Fig. 44), gonopore
basad of midlength of shaft; connective short,
not reaching apex of style (Fig. 41); style with
inner lateral margin serrate, outer one
smooth, stylar spine short, bluntly pointed
and directed laterad (Fig. 43).

Female. — Seventh sternum with caudal
margin shallowly and broadly concave, small

116

Great Basin Naturalist Memoirs

No. 12

Figs. 40-44. Colladonus claustrus (DeLong); 40, male pygofer, lateral view; 41, connective and right style, dorsal
view; 42, aedeagus, lateral view; 43, apex of right style, enlarged dorsal view; 44, apex of aedeagus and distal processes,
enlarged dorsal view.

indentation medially.

Distribution. — This species is known only
from the Mexican states of Veracruz and Chia-
pas.

Host. — Specimens were collected on un-
specified shrubs or trees at elevations be-
tween 4,000 and 7,000 feet from August to
October (DeLong 1946).

Remarks. — The female holotype and male
allotype, although from widely separate local-
ities, appear to be properly associated. From
bicinctus, to which it is closely related, claus-

trus can be separated by the absence of the
narrow, transverse, black band on the middle
of the pronotum, by the stouter pygofer spine,
by the serrated, inner lateral margin of the
stylar shaft, and by the more basad position of
the gonopore.

Colladonus verecundus (DeLong)
Figs. 4.5-50

Idiodonus verecundus DeLong 1946:20 [Holotype ?

(OSU) (examined).]
Colladonus verecundus: Nielson 1957:51
Idiodonus verecundus: Metcalf 1967:1304

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NiELSON: COLLADONUS OF MEXICO AND CENTRAL AMERICA

117

Figs. 45-50. Colladonus verecundus (DeLong); 45, male pygofer (allotype oi vereciindus), lateral view; 46, male
pygofer (allotype of acus), lateral view; 47, connective and right style, dorsal view; 48, aedeagus, lateral view; 49, apex
of right style, enlarged dorsal view; 50, apex of aedeagus and distal processes, enlarged dorsal view.

Idiodonus verecundus: DeLong 1984:11

Idiodonus acus DeLong 1946:20 [Holotype 9 (OSU) (ex-
amined).] New synonymy

Colladonus acus: Nielson 1957:51

Idiodonus acus: Metcalf 1967:1286

Idiodonus acus: DeLong 1984:11

Idiodonus mexicanus: DeLong 1946:27 [Holotype 9
(OSU) (examined).] New synonymy

Idiodonus mexicanus: Metcalf 1967:1300

Idiodonus mexicanus: DeLong 1984:11

Length. — Male, 4.5-4.6 mm; female, 4.9-
5.0 mm.

A well-marked species with two distinctive,
black spots on anterior margin of crown and
narrow to broad, ivory or yellow, transverse
band along middle of pronotum. Similar to
other pronotal banded species, but with
unique pygofer spine.

Head with anterior margin produced, apex
angled.

Male. — Pygofer in lateral view with long,

dentate caudoventral spine, spine projecting
caudodorsally with its exposed distal part
about 1/2 as long as its entire length from its
basal origin (Figs. 45, 46); aedeagus in lateral
view with bifurcated distal processes about
1/2 as long as shaft, apices reaching gonopore
(Fig. 48), crossing over in dorsal view (Fig.
50), gonopore at midlength of shaft; connec-
tive long, extending distally beyond apex of
style (Fig. 47); style with stylar shaft serrate on
inner lateral margin with short stylar spine
projecting laterally (Fig. 49).

Female. — Seventh sternum with caudal
margin slightly rounded or angled on either
side of median, shallow indentation; some-
times with slight protrusion laterally next to
indentation.

Distribution. — This species is restricted
to the Mexican states of Distrito Federal,
Michoacan, and Morelos.

118

Great Basin Naturalist Memoirs

No. 12

Figs. 51-55. CoUadonus tituhis (DeLong): 51, male pygofer, lateral view; 52, connective and right style, dorsal view;
53, aedeagus, lateral view; 54, apex of right style, enlarged dorsal view; 55, ape.x of aedeagus and distal processes,
enlarged dorsal view.

Host. — Specimens were collected on pine
at elevations from 9,500 to 10,300 feet during
September and October (DeLong 1946).

Remarks. — The female seventh sterna of
the holotypes of verecundus and mexicanus
are identical in configuration. In acus the lat-
eral margins are angled toward the middle. I
consider these traits normal variation in the
population. The sexes of each species oi vere-
cundus and acus appear to be properly associ-
ated, although there is some variation in color
between the two species. The color patterns
and configuration of the seventh sternum of
the holotype oi mexicanus are similar to vere-

cundus. The male genitalic features of the
allotypes of acus and verecundus are identi-
cal.

This species can be distinguished from all
members of the genus CoUadonus by its
unique, very long, dentate pygofer spine.

CoUadonus titulus (DeLong)
Figs. 51-55

Idkxiomi.s titulu.s DeLong 1946:24 [Holotype S (OSU)

(examined).]
CoUadonus titulus: Nielson 1957:51
Idiodonus titulus: Metcalf 1967:1302
Idiodonus titulus: DeLong 1984:11
Idiodonus goodi: DeLong 1946:27 [Holotype 9 (OSU)

1988

NiELSON: COLLADONUS OF MEXICO AND CENTRAL AMERICA

119

(examined).] New synonymy
Idiodonus '^oodi: Metcalf 1967:1298
Idiodonus goodi: DeLong 1984:11

Length. — Male, 4.4-4.6 mm; female,
4.9-5.2 mm.

General color light tannish to yellow with
deep fuscous to black markings on crown,
pronotum, and forewings; crown with two
subtriangular spots on anterior margin;
pronotum with anterior border black, broad,
transverse, tannish band below; forewings
with veins tannish to yellow, cells suffused
with light fuscous to black. Similar in general
habitus and certain male genital characters to
claustrus, but more nearly related in male
genital characters to singularius.

Head with anterior margin not produced,
apex slightly angled.

Male. — Pygofer in lateral view with long
spine arising about middle of caudal margin
and directed nearly dorsad (Fig. 51); aedeagus
in lateral view with short, rather stout, distal,
bifurcated processes not reaching midlength
of shaft (Fig. 53), crossing over in dorsal view
(Fig. 55), gonopore slightly distad of middle of
shaft; connective long, extending distad of
apex of style (Fig. 52); style with narrow stylar
shaft, stylar spine long, narrow, directed lat-
erad (Fig. 54).

Female. — Seventh sternum with caudal
margin slightly produced along middle, with
small excision at middle.

Distribution. — This species is known only
from the Mexican states of Distrito Federal
and Veracruz.

Host. — Reported from unspecified trees or
shrubs above 7,500 feet during September
and October by DeLong (1946).

Remarks. — The male holotype and female
allotype of titulus appear to be correctly asso-
ciated. The general habitus and female sev-
enth sternum of the holotype of goodi are
identical to the female allotype of titulus.
Males o( goodi are unknown.

Colladonus titulus can be separated from
claustrus by the much longer pygofer spine
and from singularius by the shorter aedeagal
processes, the more distal position of the
gonopore, and the shorter stylar spine.

Colladonus singularius, n. sp.
Figs. 56-59

Length. — Male, 4.6 mm; female, 5.0-5.2
mm.

Color tan to pale yellow with two small
spots on anterior margin of crown, narrow,
dark, transverse band on middle of crown in
male, marking incomplete in female; prono-
tum with broad, ivory, transverse band;
forewings with veins pale yellow, cells
translucent; face yellow with dark markings
on clypeus. Similar to belli in general habitus
but with distinctive male genitalia.

Head with anterior margin not produced,
apex rounded.

Male. — Pygofer in lateral view with pro-
duced caudoventral margin, pygofer spine
very long, arising from apex of caudoventral
margin and projecting caudodorsally (Fig. 56);
aedeagus in lateral view with narrow shaft and
long, distal, bifurcated processes extending
beyond midlength of shaft (Fig. 58), crossing
over in dorsal view (Fig. 57), gonopore basad
of middle of shaft; connective moderately
long, nearly reaching to apex of style (Fig. 57);
style with narrow stylar shaft, curved laterally
and with short, bluntly pointed stvlar spine
(Fig. 59).

Female. — Seventh sternum with truncate
caudal margin with small, narrow, median ex-
cision.

HoLOT\'PE (male). — Me.xico: Guanajuato,
10 mi south San Luis de la Paz 24.X. 1981, M.
W. Nielson (Nielson collection), female allo-
type, same data as holotype (Nielson collec-
tion). Paratype: seven females, same data as
holotype (BYU, Nielson collection).

Remarks. — From belli, to which it is simi-
lar in general habitus, singularius can be sepa-
rated by the very long pygofer spine and the
longer, aedeagal, bifurcated processes.

Colladonus trabilis, n. sp.

Figs. 60-63

Length. — Male, 4.8 mm.

Color. — Yellow and black. Crown yellow
with two black spots on anterior margin,
broad, black marking on either side of middle
on disc; pronotum black with broad, yellow,
longitudinal band on middle and yellow mark-
ings on lateral angles; scutellum black with
large, yellow marking medially; forewings
black with claval veins yellow except for black
marking on basal part of posterior portion,
remaining veins pale ivory, cells fuscous ex-
cept in translucent costal area, first and sec-
ond apical cells, and appendix; face black ex-
cept yellow above. Similar to tessellatus.

120

Great Basin Naturalist Memoirs

No. 12

Figs. 56-59. Colladonus singtilariiis, n. sp. : 56, male pygofer, lateral view; 57, connective, right style, and aedeagus,
dorsal view; 58, aedeagus, lateral view; 59, ape.\ of right style, dorsal view.

Head with anterior margin produced, apex
rounded.

Male. — Pygofer in lateral view with
acutely angled caudal margin, pygofer spine
moderately long and arising from apex of cau-
dal margin, spine curved mesad and directed
caudodorsad (Fig. 60); aedeagus in lateral
view with short, distal, bifurcated processes,
not reaching midlength of shaft (Fig. 62), pro-
cesses crossing over in dorsal view (Fig. 61),
gonopore at midlength of shaft; connective
long, nearly reaching to apex of style (Fig. 61);
style with broad stylar shaft, curved laterad,
outer lateral margin serrate, stylar spine very
small, blunt distally and projecting laterally
(Fig. 63).

Female. — Unknown.

HoLOTYPE (male). — Mexico: D.F., De-
sierto de los Leones, 9,600 ft, 6. VII. 1961,
George W. Byers (KU).

Remarks, — This unusual, marked species
can be distinguished from tessellatus by the
distinctive markings on the pronotum and

clavus; the broad, curved styler shaft; the
shorter, distal, bifurcated processes of the
aedeagus; and the gonopore at midlength of
the shaft.

Colladonus clathrus (DeLong)

Figs. 64-68

Idiodonus clathrus DeLong 1946:28 [Holotype $ (OSU)

(examined).]
Colladonus clathrus: Nielson 1957:51
Idiodonus clathrus: Metcalf 1967:1288
Idiodonus clathrus: DeLong 1984:11
Idiodonus turpiter DeLong 1946:28 [Holotype 9 (OSU)

(examined). ] New synonymy
Colladonus turpiter: Nielson 1957:51
Idiodonus turpiter: Metcalf 1967:1303
Idiodonus turpiter: DeLong 1984:10

Length — Male, 4.7-4.9 mm; female,
5.0-5.2 mm.

General color tawny with fuscous mark-
ings on disc of crown and forewings of mature
specimens; crown with two distinctive, black,
handlebar-shaped markings on anterior mar-
gin; pronotum without transverse band. Simi-
lar in general habitus to anademus (DeLong)

1988

NiELSON: COLLADONUS OF MEXICO AND CENTRAL AMERICA

121

Figs. 60-63. CoUadonus trabilis, n. sp.; 60, male pygofer, lateral view; 61, connective, right style, and aedeagus,
dorsal view; 62, aedeagus, lateral view; 63, apex of right style, enlarged dorsal view.

but with distinctive male genitalia.

Head with anterior margin not produced,
apex rounded.

Male. — Pygofer in lateral view with long,
robust spine arising from middle of caudal
margin and projecting caudodorsally (Fig. 64);
aedeagus in lateral view with distal, bifur-
cated processes more than 1/2 half as long as
shaft, reflexed at distal half (Fig. 66), pro-
cesses parallel in dorsal view except for distal
third (Fig. 68), gonopore distad of midlength
of shaft; connective long, extending beyond
apex of style (Fig. 65); style with narrow stylar
shaft, inner lateral margin serrate, outer one
smooth, stylar spine broad, projecting later-
ally.

Female. — Seventh sternum with broad, V-
shaped, median emargination, caudal margin
on either side convex.

Distribution. — This species appears to
be restricted to the Mexican state of Distrito

Federal.

Host. — Unknown.

Remarks. — The female holotype and male
allotype o{ clathrus appear to be properly as-
sociated. The female holotype of turpiter is
identical with the female holotype of clath-
nis , based on similar color patterns (handle-
bar-shaped markings on anterior margin of
crown) and configuration of the seventh ster-
num. Although DeLong did not designate an
allotype of turpiter in his description of that
species, a specimen so labeled was found and
examined. This specimen could not be associ-
ated with the female holotype o( turpiter but
is identical in color patterns and male genitalia
to the male allotype of nigridens, which is
treated as a junior synonym of albocinctus in
this paper.

From anademus, to which it is similar in
general habitus, clathrus can be separated

122

Great Basin Naturalist Memoirs

No. 12

Figs. 64-68. CoUadonus clathrus (DeLong): 64, male pygofer, lateral view; 65, connective and right style, dorsal
view; 66, aedeagus, lateral view; 67, apex of right style, enlarged dorsal view; 68, ape.x of aedeagus and distal processes,
enlarged dorsal view.

from all known Mexican and Central Ameri-
can species by the reflexed, distal, bifurcated
aedeagal processes.

CoUadonus beameri (Ball)
Figs. 69-75

Idiodonus beameri: Ball 1937:28 [Holotype ? (USNM)
(examined).]

Idiodonus beameri: DeLong 1946:17

CoUadonus beameri: Oman 1949:125

CoUadonus beameri: Nielson 1957:45

CoUadonus beameri: Metcalf 1967:1244

Idiodonus beamercUus: DeLong 1983:92 [Invalid re-
placement name for nonexistent species, Id-
iodonus beameri: DeLong 1946:17 nee Idiodonus
foeamert Ball 1937:28.]

Idiodonus beamereUus: DeLong 1984:10

Idiodonus nuirginatus: DeLong 1983:90 [Holotype 6

(OSU) (examined).] New synonymy
Idiodonus marginatus: DeLong 1984:10

Length. — Male, 4.2-4.6 mm; female,
5.0-5.3 mm.

General color tawny throughout with black
spots along anterior margin of crown, spots
sometimes numbering as many as eight, four
on anterior margin and two each above and
below margin next to eye, spots sometimes
connected. Similar in general habitus to in-
cidiis (DeLong) but larger and with additional
spots on crown and distinctive male genitalia.

Head with anterior margin not produced,
apex rounded.

Male. — Pygofer in lateral view with very

1988

NiELSON: COLLADONUS OF MEXICO AND CENTRAL AMERICA

123

Figs. 69-75. CoUadonus beameri (Ball): 69, male pygofer, lateral view; 70, male pygofer (holotype ofnuirginatus),
lateral view; 71, connective and right style, dorsal view; 72, aedeagus, lateral view; 73, aedeagus (holotype of
tnarginatus), lateral view; 74, apex of right style, enlarged dorsal view; 75, apex of aedeagus and distal processes,
enlarged dorsal view.

long spine, exposed length about as long as
fused basal half (Figs. 69, 70); aedeagus in
lateral view with short, bifurcated distal pro-
cesses, processes reaching to gonopore (Fig.
72, 73), crossing over in dorsal view (Fig. 75),
gonopore distad of middle of shaft; connective
long, extending to or slightly beyond apex of
style (Fig. 71); style with long stylar shaft,
lateral margins smooth, stylar spine very short
and projecting laterally (Fig. 74).

Female. — Seventh sternum with subtrun-
cate caudal margin, with shallow, V-shaped
emargination medially.

Distribution. — This is a widespread spe-
cies known from the Mexican states of Mi-
choacan, Distrito Federal, Puebla, Hidalgo,

Veracruz, Jalisco, and Morelos, and from the
state of Arizona.

Host. — Specimens were collected from
pine at elevations from 5,500 to 9,800 feet
during October in Mexico (DeLong 1946)

Remarks. — DeLong (1983) proposed heam-
erellus as a replacement name for his sup-
posed "beameri Delong" treated as beameri
Ball in his 1946 paper. That action was actually
based on a nonexistent species.

The male holotype of marginatus is identi-
cal to authentically determined males of
beameri. From incidus, to which it is similar
in general habitus, beameri can be distin-
guished by the eight spots on the anterior
margin of the crown and by the very long
pygofer spine.

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Figs. 76-79. Colladonus tessellatus , n. sp. : 76, male pygofer, lateral view; 77, connective, right style, and aedeagus,
dorsal view; 78, aedeagus, lateral view; 79, apex of right style, enlarged dorsal view.

Colladonus tessellatus, n. sp.

Figs. 76-79

Length. — Male, 4.3 mm; female, 4.7-4.9
mm.

Color tan to tannish brown with dark brown
markings on dorsum; crown with two large,
subquadrate spots on anterior margin, small
spot in ocellocular area next to lateral frontal
suture, two large spots on disc separated
widely at middle; pronotum with dark, mosaic
markings separated by narrow, tannish line
medially; forewings with dark markings in
cells, veins mostly tannish to yellow tannish;
face tan with dark markings. Similar to trahilis
in male genitalic characters.

Head with anterior margin produced, ape.x
rounded.

Male. — Pygofer in lateral view with caudal
margin obtusely angled, pygofer spine mod-
erately long, arising at apex of caudal margin
above middle and slightly curved dorsad (Fig.
76); aedeagus in lateral view narrow along

major portion of shaft, with moderately long,
distal, bifurcated process (one process broken
off in holotype) (Fig. 78) reaching to about
midlength of shaft, gonopore basad of mid-
length of shaft; connective short, not reaching
apex of style (Fig. 77); style with narrow stylar
shaft, inner lateral margin serrate, slightly ta-
pered distally and curved laterad, stylar spine
very small (Fig. 79).

Female. — Seventh sternum with truncate
caudal margin, very small, shallow excavation
medially.

Holotype (male). — Mexico: Mex. D.F., 5
mi west of Tlamacas, P.N. Ixtapopo, 10,000 ft,
31. V. 1974, C. W. and L. O'Brien and Mar-
shall (BYU), female allotype, same data as
holotype (BYU). Paratypes: one male and one
female, same data as holotype (author s collec-
tion); three females, same data as holotype
(BYU); one female, 11 mi east of Amecameca,
10,300 ft, 31. V. 1974, C. S. and L. O'Brien
and Marshall; one female, P.N., Cumbres de

1988

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125

Figs. 80-84. Colladonus anademus (DeLong): 82, male pygofer, lateral view; 81, connective and right style, dorsal
view; 82, aedeagus, lateral view; 83, apex of right style, enlarged dorsal view; 84, apex of aedeagus and distal processes,
enlarged dorsal view.

Ajusco, Llano de Cantimplora, 3,340 m,
4.IX.1982, C. W. and L. B. O'Brien and G. J.
Wibmer (CAS).

Remarks. — From trabilis, to which it is
similar in certain male genitalic features, tes-
sellatus can be separated by the gonopore,
which is basad of midlength of the aedeagal
shaft, by the short connective, and by the
slender stylar shaft, which is serrate on the
inner lateral margin.

Colladonus anademus (DeLong)
Figs. 80-84

Idiodonus anademus DeLong 1946;29 [Holotype 9

(OSU) (examined).]
Colladonus anademus: Nielson 1957:51
Idiodonus anademus: Metcalf 1967:1286
Idiodonus anademus: DeLong 1983:90
Idiodonus anademus: DeLong 1984:10

Length. — Male, 4.8 mm; female, 4.9-5.1
mm.

General color tawny throughout with two
distinct, black, elongate, triangular spots on
anterior margin of crown, small black mark-
ings on disc of crown and pronotum. Similar in
general habitus to incidus (DeLong) but with
distinctive male genitalia.

Head with anterior margin not produced,
apex rounded.

Male, — Pygofer in lateral view with cau-
doventral margin produced distally, subtrun-
cate and serrate (Fig. 80), pygofer spine very
short, basal 3/4 sclerotized and fused to inner
mesal margin of pygofer, spine projecting dor-
sad (Fig. 80); aedeagus in lateral view with
slender, distal, bifurcated processes reaching
to midlength of shaft (Fig. 82), crossing over
in dorsal view (Fig. 84), gonopore at about
middle of shaft; connective long, nearly
reaching to apex of style (Fig. 81); style with
long, narrow stylar shaft, inner lateral margin

126

Great Basin Naturalist Memoirs

No. 12

Figs. 85-89. Colladonus incidus: 85, male pygofer, lateral view; 86, connective and right style, dorsal view; 87,
aedeagus, lateral view; 88, apex of right style, enlarged lateral view; 89, apex of aedeagus and distal processes, enlarged
dorsal view.

serrate, outer one smooth, stylar spine short,
blunt, and directed laterad.

Female. — Seventh sternum with caudal
margin convex, shallow, narrow, V-shaped
excision medially.

Host. — Specimens were collected on pine
by DeLong (1946).

Distribution. — This species is known only
from the Mexican state of Distrito Federal.

Remarks. — The female holotype and male
allotype appear to be correctly associated.
This species, similar to incidus, can be distin-
guished by the produced, subtruncate cau-
doventral margin of the pygofer, by the short,
dorsally directed pygofer spine, and by the
long, narrow stylar shaft.

Colladonus incidus (DeLong), n. comb.
Figs. 85-89

Idiodonus incidus DeLong 1946:29 [Holotype 9 (OSU)

(examined).]
Idiodonus iticidus: Metcalf 1967:1299
Idiodonus incidus DeLong 1984:11
Idiodonus pallidas: DeLong 1983:90 [Holotype 6 (OSU)

(examined).] New synonymy
Idiodonus pallidus: DeLong 1984:10

Length. — Male, 4.8 mm; female, 4.9-5.1

General color tawny throughout with two
small, black spots on anterior margin of
crown. Similar in general habitus to anade-
ynus but with distinctive male genitalia.

Head with anterior margin produced, apex
angled.

Male. — Pygofer in lateral view with
broadly angled, serrate caudal margin, very
small spine at middle of margin, spine di-
rected distad (Fig. 85); aedeagus with slender,
distal, bifurcated processes extending beyond
midlength of shaft (Fig. 87), crossing over in

1988

NiELSON: COLLADONUS OF MEXICO AND CENTRAL AMERICA

127

dorsal view (Fig. 89), gonopore at midlength
of shaft; connective short, not reaching to apex
of style (Fig. 86); style with slender stylar
shaft, lateral margins smooth, stylar spine
long, directed laterad (Fig. 88).

Female. — Seventh sternum with broadly
convex caudal margin, very small indentation
medially.

Distribution — This species is known only
from the Mexican state of Distrito Federal.

Host. — Specimens were collected from
unspecified trees or shrubs at 8,500 feet dur-
ing September and October in Mexico (De-
Long 1946).

Remarks, — No males of incidus were de-
scribed in the type series. The female holo-
type of incidus was associated with the male
holotype of paUidus and is identical with the
female allotype ofpaUidiis . The latter name is
the more recent and is, therefore, suppressed
as a junior synonym oi incidus.

From anademus, to which it is similar, in-
cidus can be easily separated by the serrated,
broadly angled caudal margin of the pygofer
with its very small, distally directed spine, by
the shorter connective, and by the longer,
distal, bifurcated processes of the aedeagus.

Paracolladonus , new genus

Type-species : Idiodonus insculptus De-
Long 1946:25.

Similar to Colladonus Ball and Dolyobius
Linnavuori in general habitus but with dis-
tinctive male genitalia. General color tannish
brown in female to black in male with dark
markings on crown, pronotum, and scutel-
lum.

Head slightly wider than pronotum, ante-
rior margin broadly rounded and slightly pro-
duced medially; pronotum with posterior
margin broadly concave; scutellum as in Col-
ladonus ; clypeus broad anteriorly; venation of
forewing similar to Colladonus, inner an-
teapical open basally.

Male pygofer with large spine on caudo-
ventral margin, setose caudodorsally; aedea-
gus long, broadly recurved, somewhat
compressed dorsoventrally, with paired, re-
curved, short, distal processes, processes
sometimes with secondary process medially,
gonopore subapical on dorsal surface; connec-
tive long and narrow; style broad at basal half,
with foot-shaped apex.

Paracolladonus insculptus

(DeLong), n. comb.

Fi^s. 90-94

Idiodonus in.sctdptiis: DeLong 1946:25 [Holotype $

(OSU) (examined).]
Idiodonus insculptus: Metealf 1967:1299
Idiodonus insculptus: DeLong 1984:11

Length. — Male, 5.3 mm; female, 5.7-5.9
mm.

Long, slender species with dark markings
on crown, pronotum, and scutellum; prono-
tum black in male, light brown in female with
dark markings on anterior margin; forewing
light to dark brown with yellowish veins.

Male pygofer with very long, stout spine on
caudoventral margin, spine projecting cau-
dodorsad, numerous setae confined to cau-
dodorsal area (Fig. 90), aedeagus in lateral
view long, recurved, with distal processes
short, broad medially (Fig. 92), sometimes
with secondary process medially (Fig. 94) (if
absent, basal attachment apparent in dorsal
view), curved dorsad in lateral view, slightly
flared laterally in dorsal view (Fig. 93), shaft
somewhat flattened dorsoventrally (Fig. 94),
gonopore subapical on dorsal surface; connec-
tive long and narrow, reaching to or beyond
apex of style (Fig. 91); style with narrow ridge
or flange from inner lateral margin above
preapical lobe to apex of style in dorsal view
(Fig. 92).

Remarks. — ^This is the only known species
assigned to the genus. It is known from the
states of Distrito Federal and Morelos in Mex-
ico. DeLong (1946) collected specimens from
pine at 10,000 feet in September and Octo-
ber. A pair of specimens were collected on the
Cuernavaca road in Mexico by Ball and Stone.

Paranurenus, new genus

Type-species: Idiodonus latidens DeLong
1946:26

Similar to Colladonus in general habitus
and Nurenus in some male genital characters.
General color tawny with black markings on
head. Pronotum with ivory, transverse band,
fore wings with ivory stripe on claval suture.

Head about as wide as pronotum; crown
produced, longer medially than next to eye;
pronotum and scutellum as in Colladonus;
forewing as in Colladonus; venation partially
obscured; clypeus long; clypellus long and
narrow.

128

Great Basin Naturalist Memoirs

No. 12

Figs. 90-94. Paracolladonufi insculptus (DeLong); 90, male pygofer, lateral view; 91, connective and right style,
dorsal view; 92, aedeagus, lateral view (allotype); 93, aedeagus, enlarged dorsal view (allotype); 94, aedeagus, dorsal
view (specimen from Cuernavaca Road, Mex.).

Male pygofer without caudal spine, margin
obliquely truncate as in Niirenus; aedeagus
small, recurved, flattened dorsoventrally with
pair of short, acuminate distal processes;
gonopore subapical on dorsal surface; connec-
tive and style as in Colladonus.

Paranurenus is represented by a single
known species from Mexico.

From Colladonus, to which it is similar in
general habitus, Paranurenus can easily be
differentiated by the flattened aedeagus and
by the absence of a pygofer spine. The head
characters and the flattened shaft of the aedea-
gus will differentiate the genus from Nurenus.

Paranurenus latidens (DeLong), n. comb.
Figs. 95-98

Idiodonus latidens: DeLong 1945:26 [Holotype 9 (OSU)
(examined).]

Idiodonus latidens: Metcalf 1967:1300
Idiodonus latidens DeLong 1984:11

Length. — Male, 4.4-4.5 mm; female, 4.8-
5.1 mm.

Color tawny, crown ivory with two black
spots on anterior margin and narrow, black,
transverse band on disc between eyes; prono-
tum tawny with narrow, ivory, transverse
band above middle; scutellum tawny; fore-
wing tawny with ivory on claval suture ex-
tending to pronotal band, fuscous band above
costal area; face ivory to tawny with black
markings.

Male. — Pygofer in lateral view with
obliciuely truncate caudal margin, pygofer
spine absent, macrosetae on caudodorsal
quadrant (Fig. 95); aedeagus in lateral view
recurved, socle large, shaft slender, flattened

1988

NiELSON; COLLADONUS OF MEXICO AND CeNTKAL AMERICA

129

Figs. 95-98. Paranurenus latidens (DeLong): 95, male pygofer, lateral view; 96, connective and right style, dorsal
view; 97, aedeagus, lateral view; 98, aedeagus, ventral view.

dorsoventrally (Fig. 97) with pair of short,
acuminate distal processes, processes di-
rected laterad in ventral view (Fig. 98); gono-
pore subapical on dorsal surface of shaft; con-
nective long and narrow, extending just be-
yond apex of style (Fig. 96); style with stylar
shaft short and enlarged distally, stylar spine
prominent, subapical, and projecting laterally
(Fig. 96).

Female. — Seventh sternum with short,
median, spatulate process on caudal margin.

Remarks. — This species has no known
close relatives. The styles are typical of many
species oi Colladonus, and the aedeagus, ex-
cept for its flattened condition, is similar in
configuration to Nurenus. The head charac-
ters and general habitus are similar to species
oiColladonus.

The species is known from the states of
Michoacan and Distrito Federal in Mexico,

and occur at elevations from 5,000 to 8,000
feet in February and September on unspeci-
fied shrubs or trees (DeLong 1946).

Jaacunga, new genus

Type-species: Idiodonus vinculus DeLong
1945:15

Similar to Paracolladonus in general
habitus but more robust with slightly wider
head and distinctive male genitalia. General
color tawny with markings on crown, narrow
band on pronotum and iridescent forewings.

Head wider than pronotum, anterior mar-
gin broadly rounded and slightly produced as
in Paracolladonus; pronotum large, posterior
margin broadly and shallowly concave; fore-
wing venation as in Paracolladonus.

Male pygofer without caudal spine, setose
submarginally; aedeagus stout with enlarged
socle, recurved at apical third with terminal

130

Great Basin Naturalist Memoirs

No. 12

Figs. 99-101. Jaacunga vinculus (DeLong): 99, male pygofer, lateral view; 100, connective, right style, and
aedeagus, dorsal view; 101, aedeagus, lateral view.

processes, gonopore large, subapical on dor-
sal surface; connective Y-shaped; style with-
out apical spine. Female seventh sternum
with median spatulate process. Two species,
described below, are assigned to the genus.

Jaacunga vinculus (DeLong), n. comb.
Figs. 99-101

Idiodonus vinculus: DeLong 1946:15 [Holotype 9 (OSU)

(examined).]
Idiodonus vinculus: Metcalf 1967:1304
Idiodonus vinculus DeLong, 1984:10

Length. — Male, 5.8 mm; female, 6.4 mm.

Long, robust species with two spots on
anterior margin of crown and narrow, black,
transverse, irregular band on disc in male,
less marked in female; pronotum with narrow,
yellow or ivory, transverse band near middle
and similar, less pronounced band on poste-
rior margin; forewing iridescent, veins black

to tawny. Similar to spatulatus in male genital
character.

Male. — Pygofer without caudal spine,
macrosetae on submargin from dorsal to ven-
tral margin (Fig. 99); aedeagus in lateral view
stout, socle enlarged, shaft tubular, curved
dorsad at apical half with six terminal spines,
two long and projecting anteroventrally, two
moderately long and projecting anterodorsad,
and two very short and projecting anteriorly
(Fig. 101), gonopore large, subapical on dorsal
surface; connective stout, Y-shaped, extend-
ing slightly beyond apex of style (Fig. 100);
style similar to CoUadonus but broader at
basal two-thirds, stylar apex narrow, without
distinct stylar spine, apex obliquely truncate.

Female. — Seventh sternum with short,
apically bifid, medial projection on caudal
margin, shallowly excavated on either side of
projection.

1988

NielsonColladonus OF Mexico AND Central America

131

Figs. 102-104. jaacunga spatiilatiis (DeLong): 102, male pygofer, lateral view; 103, connective, right style, and
aedeagus, dorsal view; 104, aedeagus, lateral view.

Remarks. — This species is known from the
female holotype and male allotype specimens
collected at Jacala, Hidalgo, Mexico, and Aca-
pulco Road, Mexico, respectively. Other
specimens that I have not seen are reported
from Mexico City and Jalapa Road in Mexico
by DeLong (1946). All specimens were col-
lected from September to October at eleva-
tions from 5,000 to 8,500 feet from unspeci-
fied shrubs or trees.

Jaacunga vinculus can be distinguished
from spatulatus by its six terminal spines,
which differ in their configuration, and by the
shape of the apex of the style.

Jaacunga spatulatus (DeLong), n. comb.
Figs. 102-104

Idiodonus spatulatus DeLong 1946:15 [Holotype 9

(OSU) (examined).]
Idiodonus spatulatus: Metcalf 1967:1302
Idiodonus spatulatus DeLong 1984:10
Idiodonus ruhellus DeLong 1946:15 [Holotype 9 (OSU)

(examined).] New synonymy
Idiodonus ruhellus: Metcalf 1967:1301
Idiodonus ruhellus DeLong 1984:9

Length. — Male, 5.5 mm; female, 5.5-5.8
mm.

Similar in general habitus to vinculus but
with distinctive male genitalia.

Male pygofer without caudal spine, mac-
rosetae submarginal in caudodorsal area (Fig.
102); aedeagus in lateral view stout but not as
robust as in vinculus, socle enlarged, shaft
similar to vinculus, with four terminal pro-
cesses, longest pair projecting ventrally and
broad before gradually pointed apex, second
pair shorter, projecting anteriorly with tips
curved ventrally (Figs. 103, 104); gonopore
large, subapical on dorsal surface; connective
similar to vinculus except shorter, not reach-
ing apex of style (Fig. 103); style as in vinculus
except apex curved mesally and rounded dis-
tally.

132

Great Basin Naturalist Memoirs

No. 12

Remarks — Jaacunga spatulatus is known
from the type series collected from Mexico
City and from the female holotype o^nibeUus,
which has the same type locality as spatulatus.
The specimens were collected from unspeci-
fied shrubs at elevations of 7,500 to 8,500 feet
in September (DeLong 1946). The species can
be distinguished from vinculus by the shape of
four terminal spines on the aedeagus and by
the shape of the stylar apex.

Paracrassana, new genus

Type-species: Idiodonus nigrifrons De-
Long 1983:91

Near Crassana DeLong & Heishberger but
with distinctive head, facial, and genitalic fea-
tures. General color tawny with black mark-
ings on head and thorax, face black.

Head slightly wider than pronotum, ante-
rior margin slightly angled and produced me-
dially, not rounded to face, clearly demar-
cated but not carinate, narrow, transverse
impression submarginally between ocelli,
surface shiny, striate longitudinally; ocellus
remote from eye; forewing as in Crassana;
clypeus broad anteriorly, about as wide as
long; clypellus long, lateral margins expanded
distally.

Male pygofer long, with deep incision
basally to near middle, dorsal margin broadly
excavated near middle, row of fine teeth on
ventral margin; aedeagus symmetrical, long,
narrow, nearly tubelike, socle basal, gono-
pore terminal, sclerotized endotheca basad of
aedeagal shaft; connective very short as in
Crassana and Neocrassana Linnavuori; plate
long, triangular, tenth segment sclerotized
laterally.

Paracrassana is represented by a single
known species from Mexico. It can be distin-
guished from Crassana by the narrow demar-
cation between the crown and face, the nar-
row impression submarginally on the crown,
the long pygofer with basal incision and tubu-
lar aedeagus. From Neocrassana, it can be
distinguished by the remote position of the
ocelli from eyes, the position of the socle
on the aedeagus, and characters described
above.

Paracrassana nigrifrons (DeLong), n. comb.
Figs. 105-107

Idiodonus nigrifrons DeLong 1983;91 [Holotype 6
(OSU) (examined).]

Length. — Male, 5.4 mm.

Long, robust species, crown with irregular,
narrow, black, transverse band between
ocelli; pronotum with black markings on ante-
rior lateral angles; face black except for nar-
row, yellow anterior margin; forewing iri-
descent, veins fuscous.

Male pygofer in lateral view long and nar-
row, macrosetae on distal third except caudal
margin, ventral margin with fine spines (Fig.
105); aedeagus long, tubular, curved dorsally
at distal third, with subdistal flange laterally
on dorsal margin (Fig. 106), gonopore apical,
endotheca not forming a loop; connective
short, broad; style narrow, with narrowed
curved apices.

Female. — Unknown.

Remarks. — This species is known only from
the male holotype from Huanchinango, Pue-
bla, Mexico. The host is unknown.

The nomenclature of a few species de-
scribed in Idiodonus and Colladonus are
treated below:

Bonneyana caldwelli (DeLong), n. comb.

/(/io(/onusca/duJe//i DeLong 1946:16 [Holotype 9 (OSU)

(examined).]
Idiodonus caldwelli: Metcalf 1967:1288
Idiodonus caldwelli DeLong, 1984:10
Idiodonus apertus DeLong 1946:14 [Holotype 9 (OSU)

(examined).] New synonymy.
Idiodonus apertus: Metcalf 1967:1287
Idiodonus apertus Delong 1984:10

I have compared dissected specimens of
caldwelli and schwartzi (Ball) and found them
to be distinct. Two remaining known species
in the genus, osborni (Ball) and terminalis
(Ball), have not been examined; but, based on
the original descriptions, they appear to be
distinct from caldwelli. Further study of the
group is warranted, however, including illus-
trations and descriptions of the male diagnos-
tic characters.

Ollarianus tripartitus DeLong

Ollarianus tripartitus DeLong 1944:398
/r/tO(/oni/sfl/fo(frons DeLong 1983:89 [Holotype 9 (OSU)

(examined).] New synonymy
Idiodonus alhifrons DeLong 1984:10

Examination of the female holotype of /.
alhifrons and comparison with specimens of
several species o{ Ollarianus show that it be-
longs in that genus and is identical with O.
tripartitus. The genus Ollarianus has wide

1988

NiELsoN: CoLLADONus OF Mexico AND Centiul America

133

107

Figs. 105-107. Paracrassana nigrifrons (DeLong): 105, male pygofer, lateral view; 106, connective, right style, and
aedeagus, dorsal view; 107, aedeagus, lateral view.

distribution from the Nearctic to the Neotrop-
ical region and needs revision.

Paratanus costatus (DeLong), n. comb.

Idiodonus costatus DeLong 1984:92 [Holotype 3 (OSU)
(examined).]

This species is close to Paratanus wygod-
zinskyi recurvatus (Linnavuori) but differs
from it in color characters. It may be a distinct
subspecies of the nominate form or a separate
species pending studies of the genus Para-
tanus.

Angulanus DeLong, n. status

Idiodonus subgenus Angulanus DeLong 1946:30. Type-
species, Idiodonus incisurtis DeLong 1946:30.
[Holotype 9 (OSU) (examined).]

Idiodonus (Angulanus): Metcalf 1967:1304

Idiodonus: DeLong 1984:9. [Inadvertent synonymy of
Angulanus DeLong in the treatment of the type-
species, incisurus DeLong, in a key to the species
oi Idiodonus .]

Examination of the genital structures of the
male allotype clearly showed that Angulanus
deserves generic-level rank. The aedeagus is
asymmetrical, and the pygofer possesses a
prominent spine on the caudal margin, char-
acters that are not descriptive of the genus
Idiodonus.

Angulanus incisurus (DeLong), n. comb.

/diodonus incisurus DeLong 1946:30 [Holotype 9 (OSU)

(examined).]
Idiodonus incisurus: DeLong 1946:13, 14
Idiodonus (Angidanus) incisurus: Metcalf 1967:1304
Idiodonus (Angulanus) incisurus: DeLong 1984:10

134

Great Basin Naturalist Memoirs

No. 12

Idiodonus plummeri DeLong

Idiodonus plummeri DeLong 1946:25 [Holotype 9

(OSU) (examined).]
Idiodonus plummeri: DeLong 1984:11
Idiodonus bakeri. DeLong 1984:26 [Holotype 9 (OSU)

(examined).] New synonymy
Idiodonus bakeri: DeLong 1984:11

The generic placement of this species ap-
pears to be correct. No males were described
in the type series oi plummeri or bakeri, but a
male paratype specimen labeled "Idiodonus
bakeri DeLong, MB 73, Mexico, A. Dampf
coll." was found and examined. The genitalic
structures clearly show that it belongs in Id-
iodonus . The species was associated with the
female holotype of bakeri, which was found to
be identical with the female holotype o( plum-
meri.

Species incertae sedis

Three species of Idiodonus described by
DeLong, edentulus, excavatus, and copidus,
are treated herein as incertae sedis until males
are known and associated with females of
these species. Idiodonus copidus appears to
be close to Jaacunga and may belong in that
genus.

Acknowledgments

I thank Dr. Charles A. Triplehorn and
Dr. Paul Cwikla, Ohio State University
(OSU), for placing on loan nearly the entire
type series of all the Idiodonus species from
Mexico and Central America described by the
late Dr. D. M. DeLong. Without this material
it would not have been possible to unravel the
taxonomic problems involved with the Id-
iodonus and Colladonus fauna in that part of
the world. I also thank Dr. James P. Kramer,
U.S. National Museum, for the loan of the
type of Colladonus delongi Linnavuori and
material of genera related to Colladonus.

My appreciation is extended to Jeanette
Price and Pamela Scott for the illustrations
and to Kaye Thome for identifying the host
plant of a population of the Colladonus belli
complex. I thank Dr. Leon Hepner for read-
ing the manuscript and offering helpful sug-
gestions to improve its content.

References Cited

Ball, E D 1937. Some new species of leafhoppers in
groups recently segregated from Thamnotettix.
Bull. Brooklyn Ent. Soc. 32: 26-31.

Bliven, B P 1954. New Hemiptera from redwood (Miri-
dae, Cicadellidae). Bull. Brooklyn Ent. Soc. 49:
119.

DeLong, D M 1944. The genus Ollarianus (Homoptera:
Cicadellidae) in North America, including Mex-
ico. J. Wash. Acad. Sci. 34: 391-398.

1946. The Mexican species oi Idiodonus (Homop-

tera-Cicadellidae). Ohio J. Sci. 46: 13-30.

1983. New species and distribution notes of Mexi-
can and Bolivian Idiodonus (Homoptera: Cicadell-
idae) of Mexico and Bolivia. Entomol. News 95(1):
9-15.

1984. A revised key to the Idiodonus (Homoptera:

Cicadellidae) of Mexico and Bolivia. Entomol.
News 95(1): 9-15.

Gillette, C. P, AND C. F. Baker 1895. A preliminary list
of the Hemiptera of Colorado. Colorado Agric.
Expt. Sta. Bull. 31. 1.37 pp.

Medler, J T 1943. The leafhoppers of Minnesota. Ho-
moptera: Cicadellidae. Minnesota Agric. Expt.
Sta. Tech. Bull. 155. 196 pp.

LiNNAVUORL R 19.59. Revision of the Neotropical Delto-
cephalinae and some related subfamilies (Ho-
moptera). Suomal. Elanja Kasv. Seuran Vanamon
Elaint. Julk. 20: 1-370.

Nielson, M W 19.57. A revision of the genus Colladonus
(Homoptera, Cicadellidae). U.S. Dept. Agric.
Tech. Bull. 11,56: 1-.52,

1962. New species of leafhoppers in the genus

Colladonus (Homoptera, Cicadellidae). Ann. En-
tomol. Soc. Amer. .55: 143-147.

1966. A synopsis of the genus Colladonus (Ho-
moptera: Cicadellidae). J. Kansas Entomol. Soc.
.39: .3.33-.336.

Oman, P W 1949. The Nearctic leallioppers (Ho-
moptera: Cicadellidae). A generic classification
and check list. Washington Entomol. Soc. Mem.
3: 1-2,53.

Stal, C 1864. Hemiptera Mexicana enumeravit spe-
ciesque novas descripsit. Stettiner Entomol. Zeit.
25: 49-86.

Uhler, P R 1877. Report upon the insects collected by
P. R. Uhler during the explorations of 1875, in-
cluding monographs of the families Cynidae and
Saldae and the Hemiptera collected by A. S. Pack-
ard, Jr. U.S. Geol. and Geog. Survey of the Terr.
Bull. 3(2): 355-475.

NEW WORLD FULGORIDAE, PART I: GENERA WITH ELONGATE

HEAD PROCESSES

Lois B. OBi-it-n'

Abstract. — Genera new to science described below include: Amerzanna, Sinuala, and Staluhra. New species
include: Amerzanna peruana (Peru), Amijcle hrevis (Mexico), A. grandis (Mexico), A. mankimi (Honduras), Aphro-
disias shaman (Mexico), Enchophora maculata and E. uniformis (Peru), Phrictus delicatiis (Brazil), P. dilinen.s
(Colombia), Sinuala schmidti (Belize), S. stall (Honduras and El Salvador), S. tuherculata (Costa Rica), and Staluhra
brunnea (Brazil and Guyana). In addition, Artacie dufourii (Signoret) is removed from synonymy with A. haemoptcra
(Perty). New generic synonymy includes C/H/oi>ifl Stal(= Ecuadoria) Distant, En/iyr/na (Walker) (= UluhraStdl). New
synonymy of species includes: Copidocephala merula Distant (= Coanaco melanoptera Schmidt), Copiducephala
viridiguttata Stal (= Coanaco ornanda Distant), Diareusa conspersa Schmidt (= D. dahli Ossiannilsson), Enchophora
nigromactdata Distant (= E. nigrolimhata Lallemand), Enchophora recurva (Olivier) (= £. hohemani Stal), Kri-
chophora sanguinea Distant (= £. /Zorens Distant and E. longirostris Distant), Enchophora tuberctdatu (Ohvit'r) (= E.
parvipennis Walker), Enchophora viridipennis Spinola (= E. emincnta Schmidt), Enhydria tessellata (Walker) (= £.
brachialis Stal), Fulgora graciliceps Blanchard (= Laternaria orthocephahi Fonseca), Fulgora laternaria (Linnaeus)
(= F. servillei Spinola), Phrictus auromactdatus Distant (= P. notatus Lellemand), Phrictus moebiusi Schmidt (= P.
sordidus Caldwell). The two species of Fulgora are synonymized by Ridout. New combinations include Chilobia
dichopteroides (Distant) (Ecuadoria) and Stahtbra ruftda (Lallemand) (Enhydria).

Although the genera of New World Fulgori-
dae were monographed in an excellent paper
by Stal (1870b), 25 of the 64 genera have been
described since then. Also, there have been
many subsequent species described, many in
the wrong genus, and there are few keys to
species. Consequently, even though they are
large and beautiful insects, it is very difficult
to identify most species. The purpose of this
paper is to provide a guide for identification
with emphasis on the use of external charac-
ters.

Very little is known about the biology of
Fulgoridae. Two species have been reported
to be of economic importance (Wilson and
O'Brien 1987), Phrictus diadema (L.) on cocoa
trees {Theobroma cacao L.) in Brazil (Silva
1945) and Pyrops candelaria (L.) on longan
and mango (Kershaw and Kirkaldy 1910).
Host records for species contained in this pa-
per include Amycle pinyonae Knull & Knull
on pinyon pine, Pinus monophylla Voss (Knull
and Knull 1947), Fulgora laternaria (L.) on
Hymenaea courbaril L. or quapinol (Janzen
1983), Enchophora pallidipunctata Lalle-
mand on Copaifera dourajeauni (Legumi-
nosae) (label data, species not found in botani-
cal references), and Rhabdocephala brunnea

Van Duzee on Baccharis sarothroides Gray.
Johnson and Foster (1986) explored host
specificity in a tropical forest, where plant
diversity might make host finding difficult,
and found 71% of Enchophora longirostris
Distant (= sanguinea Distant) on Simarouba
amara Aubl. (Sapindales) and 84% of Phric-
tus quinquepartitus Distant on Terminalia
oblonga Steud. (Combretaceae). Adults have
been reported to aggregate on tree trunks in
ranks and to move in conjunction with each
other (Johnson and Foster 1986, O'Brien and
Wilson, 1985). Further host data will be given
in parts to be published later.

Little information on life histories is avail-
able. Kershaw and Kirkaldy (1910) noted that
the eggs o{ Pyrops candelaria were laid in flat
clumps on tree trunks and covered with wax
from the female abdominal segments; nymphs
dispersed and fed on branches; and adults fed
on tree trunks. Hingston (1932) reported rear-
ing fulgorid nymphs from what he described
as a mantidlike egg case. Other descriptions of
immatures are those of Fulgora phosphorea
L. (= laternaria L.) (Hagmann 1928); Pris-
tiopsis serratus (F.) (= Cathedra serrata [¥.])
(da Fonseca 1931), and Itzalana subnuiculata
Schmidt (Wilson and O'Brien 1986). As far as

'Entomology, Florida A & M University. Tallahassee. Florida, USA.

135

136

Great Basin Naturalist Memoirs

No. 12

is known, fulgorids feed in phloem tissue and
are univoltine. No accounts of other typical
fulgoroid behaviors, such as accoustical com-
munication and association with ants, have
been reported.

Fulgoridae are subject to predation by fly-
catchers (D. Wechsler, personal communica-
tion) and parasitization by the epipyropid
moths (Lepidoptera, Epipyropidae). The col-
ors of the closed wings would appear to make
them inconspicuous against tree trunks, but
they can be found fairly easily by the human
eye if observed in profile. I assume that the
red, orange, yellow, and white coloration so
common at the base of the hind wings is simi-
lar in function to that of the underwing moths
(Noctuidae), in which the color has been
shown to startle birds seeking prey. Eye spots
on the hind wings of Cathedra and Fulgora
would seem to serve a startle or warning func-
tion as well. But for all the speculation on the
bizarre shapes of the head processes, no one
has reported the response of predators to the
"alligator head" of the peanut bug, nor to any
other species.

One common name for the family Fulgori-
dae, "lantern-flies," was given because Ful-
gora were reported to emit light (de Merian
1771), but recent inquiries and enzyme stud-
ies have failed to substantiate this phe-
nomenon (Ridout 1983). Many Amerindians
feared Fulgora, reporting that they flew a zig-
zag path through the forest, killing everything
they touched (Branner 1885). Shamans car-
ried one in their amulet bags. The modern
version of this myth, probably invented and
surely repeated in coed entomology classes far
out in the bush, is that if a human is bitten by a
Fulgora, death is certain unless one is saved
by the antidote, mating within 24 hours; non-
sense, of course.

Lallemand (1959, 1963) revised the 20 gen-
era and 102 species of Fulgoridae from Africa
and the 28 genera and 178 species of Asia and
Australia. The New World has a much richer
fauna, composed of 64 genera and 250 species
as I begin this monograph. The only revisions
of New World genera in which authors ex-
amined types are Cyrpoptus (Kramer 1978)
and Fulgora (Ridout, personal communica-
tion). Other generic reviews are those on Ful-
gora (da Fonseca 1932), Enchophora (Metcalf
1938), Phrictus (Caldwell 1945), and Poiocera
(Gerstaecker 1860), which are either out of

date or relied on published descriptions. I
have examined types of all but the following
species: Artacie dufourii (Signoret) (MZF?,
letter not answered); A. haeriioptera (Perty)
(ZSBS, not found); Cathedra serrata (Fabri-
cius) (not listed by Zimsen 1964); Copidoced-
phala melanoptera Schmidt, Diareusa con-
spersa Schmidt, and Enchophora etninenta
Schmidt (IZW); Enchophora ensifera (Ger-
mar) and E. tuba (Germar) (lost in Lvov, Rus-
sia); E. nigrolimbata Lallemand and Enhydria
rufula Lallemand (FSAG, sent but not yet
arrived); Olivier s species, £. recurva, E. tu-
berculata, and F. caerulescens, and the types
of Fulgora, which were studied by Ridout.
Olivier's (1791) species were redescriptions
with Latin binomials of Stoll's (1781, 1788)
descriptions and figures in French and Dutch
or Flemish, using names in those languages.
Olivier's species are not in the Paris Museum
(MHNP), nor in his personal collection de-
posited there, which is all Coleoptera (Bour-
goin, personal communication). Olivier cites
"Du cabinet de M. Holthuisen" in some spe-
cies; Horn and Kahle (1935) say of Holthuisen
collection "in Hamburg am 3. II. 1796 u.
16. V. 1797 sowie in Stralsund 1800 verauk-
tionert." If any specimens went to the Ham-
burg Museum (ZMUH), they were destroyed
by bombing in World War II, except for a few
specimens out on loan. I have not been able to
trace Olivier's or Stoll's specimens further.
The first three of the species listed here were
well illustrated when described; Schmidt's
descriptions clearly fit the specimens I have;
only Lallemands, Germar's, and Olivier's
species are in doubt.

Fulgoridae may be distinguished from
other Fulgoroidea by the presence of numer-
ous cross-veins in the hind wings, a feature
lacking in all other families. Stal also noted
that the carina between the frons and gena
continues onto the clypeus, but this feature
also occurs in some Derbidae, Dictyophari-
dae, and Lophopidae. Fulgoridae share with
the Dictyopharidae similar bilaterally sym-
metrical, usually trilobed, inflatable male
genitalia. Emelyanov (1979) transferred some
Asian genera from Dictyopharidae to Ful-
goridae, citing other characters, but these
characters are not diagnostic for New World
families.

The stable characters easiest to use are
color, especially of the hind wing; the shape

1988

O'BRIEN; New World Fulgoridae

137

and length of the head process when present;
and the shape and carinae of the frons, vertex,
and thorax. Since the proportion of the head
length to the body length varies greatly with
the genera and cannot be determined from
the illustrations, measurements are given for
the genera (unless it is impossible to get an
accurate measurement because of the vari-
ability of the curvature of the head process as
in Enchophora). The ratio of head/pronotum
is given for most species. Male genitalia are
difficult to inflate consistently, but they are
figured for some specimens; I state when I
believe they must be used for a valid determi-
nation, such as in Scolopsella. These descrip-
tions are brief and diagnostic; illustrations are
more helpful than words. A character matrix is
being prepared and will be published when all
of the genera have been studied.

Part I covers those genera with head pro-
cesses. These keys are artificial. When the
definition of genera and placement of species
is completed, an attempt will be made to pro-
duce cladograms and to identify monophyletic
taxa. Two species are consigned to incertae
sedis. I do not recognize the genus of En-
chophora ensifera (Germar) among any New
World Fulgoridae or Dictyopharidae, nor
Fulgora caerulescens Olivier as figured by
Stoll (1788, plate 13, Fig. 65) among any New
World Fulgoridae or Flatidae.

Illustrations of the characters used are
found in Figures 1-10 and 33. Three words
that may not be commonly used elsewhere are
defined as ioWows: fossette — a small, deep pit;
porrect — extending forward horizontally;
terete — round in cross section. The head pro-
cess may have 10 carinae or fewer, which are
named in Figure 4. The median vein is pecti-
nate in Figure 5. The convention in Ful-
goroidea is to consider the "base" of the insect
the dorsal junction between head and thorax,
with the base of the head and pronotum both
here. The ventral part of the head has its base
anterad, at the junction with the vertex, and
its apex at the frontoclypeal suture, which is
the base of the clypeus.

To save duplication in each section, 1 have
included within the final part of this paper the
literature cited for all parts (although papers
published subsequent to Metcalf 's catalog are
cited here); a fist of localities with provinces or
longitude and latitude (only the city or dis-
tance for a city plus elevation where given will

be included earlier), and leclotypes or neo-
types will be designated. There is a list of
genera and species and their distribution by
country at the end of this part. Type reposi-
tory abbreviations are listed in the acknowl-
edgments.

To complete this monograph successfully, I
will need help in locating specimens, since
many species are known from uni(}ues or one
sex only, and distribution records are incredi-
bly incomplete. For at least a year after the
publication of this article, I will accept speci-
mens for identification for the traditional rec-
ompense, a sample of the specimens in return
for identification. Perhaps your specimens
will become types. The genus Chilohia is a
case in point: there are four described species,
1 have 10 new ones on hand, and only one
species is known from more than one speci-
men. Please help if you have unidentified
specimens.

Key to Genera of New World Fulgoridae
with Head Processes

1 . Tegmen usually transparent, rarely opaque in

basal half 2

— Tegmen opaque, rarely transparent at apex . . 4

2(1). Tegmen transparent brown with small bris-
tles on veins; prothorax with small, white nod-
ules Enhydria

— Tegmen without bristles on veins; prothorax
without small, white nodules 3

3(2). Vertex longer than broad (Fig. 1) Stalubra

— Vertex 3 times as broad as long in midline,
emarginate medially (Figs. 51, 54, 60) or apex
of vertex hidden by recurved head process

(Fig. 57) Chilohia

4(1). Each hind wing with an eyespot visible from
below with wings unspread; head process gib-
bous or laterally serrate with narrow apex;
large species, over 50 mm in length 5

— Hind wings without large, round eyespots;
head process porrect or curved, but not as
above 6

5(4). Head process peanut-shaped, in lateral view
with markings mimicking an alligator head;
eyespot on hind wing with iris or pupil Fulgora

— Head process subtriangular, narrowing at
apex, strongly serrate (Fig. 26); eyespot on
hind wing without iris or pupil Cathedra

6(4). Head process porrect (Figs. 14-32, 46-50,

102-117) 9

— Head process curved up and back (Figs.
33-35, 40-42, 63-76) 7

7(6). Preocular horizontal flange between eye and
junction of frons and vertex; head process ad-
pressed to vertex, flattened, rugosely tuber-
culate (Figs. 33-35) Artacie

138

Great Basin Naturalist Memoirs

No. 12

Figs. 1-13. See facing page for identification.

1988

O'BRIEN; New World Fulgoridae

139

Figs. 1-13. 1-3, 5, 6: Staluhra hrevis O'Brien: 1, dorsal view, head and thorax; 2, frons and clypeus; 3, lateral view,
head; 5, tegmen; 6, hind wing. Hypothetical insect: 4, diagramatic cross section of head process showing position of
carinae; 7, stylized hind tibia. Male genitalia, lateral, dorsal, and ventral views: 8-10, Scolopsella mexicana O'Brien;
11-13, Scolopsella reticulata Ball.

Morphological characters:

a-antenna

A,-lst anal vein

aed— aedeagiis

af-anal fold of hind wing

all- anal flap (10th and 1 1th abdominal segments)

an— anal area

ant— anterior margin of wing

ap— apical area of hind wing

ax-apex

b-base

C— costa or costal vein

c— clypeus

cl-clavus

clas-clasper

cm— commissural margin

cor-corium

cs— claval suture

CU— cubitus or cubital vein

cx-coxa

e-eye

f-frons

fe— femur

fo-fossette

g-gena

hp-head process

Ic^lateral carina

Ih-length of head

Ihp-length of head process

Ip-length of pronotum

M -media, medial vein

m-mesonotum

mac-marginal carina

mc-median carina

mem-membrane of tegmen

nl-nodal line

o-ocellus

p-pronotum

pc-pleural carina

Pcu-postcuhital vein

pf-postocular flange

post-posterior area of wing

pr-pronotum

prf-preocular flange (see Fig. 33)

py-pygofer

R-radius, radial vein

sa— sutural angle

Sc-subcostal vein

sf-supraocular flange

t-tegula

ta-tarsomeres

ti-tibia

tr-trochanters

vertex of head

(Continued from page 137)

— Preocular horizontal flange absent; head process
usually not adpressed to vertex (see E. tuber-
culata, subviridis, pyrrhocrifpta. Figs. 67,
73, 75) ' 8

8(7). Pronotal median carina raised, with deep fos-
settes at base; mesonotal apex depressed, stri-
ate (Figs. 63-76) Enchophora

— Pro- and mesonota smooth and gently
rounded (Figs. 40-42) Copidocephala

9(6). Medium-sized species, 26-50 mm in length

— Small species, under 25 mm in length 13

10(9). Head process with apex twice as wide as nar-
rowest part, dorsolateral edges of process
with spines Phrictus

— Head process with apex not so widened, sides

of process without spines 11

11(10). Sutural angle of tegmen triangularly pro-
duced; head with small, horizontal, preocular
flange (Figs. 116-117) Odontoptera

— Sutural angle of tegmen rounded; head with
slight vertical flange on vertical carina (Figs.
27-32) 12

12(11). Head process elongate, head 1/3 length of
insect, subtriangular, slightly bent dorsad at
apex; hind wing clear with veins tinged with
green at base, brown elsewhere .... Ainerzanna

— Head process proportionally shorter, less
than 1/5 length of insect, with sides subparal-
lel, apex slightly enlarged, not bent dorsad;
hind wing brown with either red or white
spots Diareusa

13(9). Head process semicircular in cross section;
with dorsal surface slightly compressed, ven-
tral rounded Amycle

— Head process more nearly rectangular or cir-
cular in cross section 14

14(13). Head, including process, longer than thorax;

tegmen with major veins straight 15

— Head, including process, shorter than thorax;
tegmen with major veins irregular, slightly
angulate 16

15(14). Head process subrectangular in cross section,
carinae foliaceous, angulate (Figs. 24, 25) . . .
Scolopsella

— Head process smoothly rounded, almost cir-
cular in cross section, carinae normal (Fig. 23)
Rhabdocephala

16(14). Costal margin of tegmen markedly sinuate
(Fig. 43); head process not transversely striate
Sinuala

— Costal margin of tegmen slightly convex; head

process transversely striate (Fig. 49)

Aphrodisias

140

Great Basin Naturalist Memoirs

No. 12

Figs. 14-28. Head and thorax, dorsal view: 14, Amycle brevis O'Brien, 15, A. pinyonae KnuW & Knull; 16, A. saxatilis
Van Duzee; 17, A. tumacacoriae Knull & Knull; 18, A. vernalis Manee; 19, A. g,randis O'Brien; 20, A. sodalis Stal; 21,
A. manhinsi O'Brien; 22, A. amabilis (Westwood); 2.3, RhahdoccpJuila hninnea Van Dnzee; 24, Scolopsella mexicana
O'Brien; 25, S. reticulata Ball; 26, Cathedra scrrata (Fahricius); 27, 28, Amcrztinna prrnana O'Brien (27, lateral view).

Amerzanna, n. gen.

Figs. 27, 28

Type-species: Amerzanna peruana O Brien.

Medium-sized, narrow, brown insect.s,
31-41 mm long, head 1/3 length of insect.

Head process porrect, terete, gradually nar-
rowed to apex; apex bent upward; dorsal and
ventral median and lateral carinae present
on anterior third; weak hut traceable pleu-
ral carina cvuving ventrally to end in slight

1988

O'BRIEN: New World Fulgoridae

141

ventral tubercle. Preocular flange absent.
Pronotum smoothly rounded, traces of fos-
settes and pleural carinae faintly visible. Teg-
men with M pectinate; cross-veins irregular,
forming diamond-shaped cells. Profemur Ion-
longer than metafemur, posterior tibia with 6
lateral spines. Ninth abdominal tergite in fe-
males twice as long as eighth.

I name this genus Amerzanna because su-
perficially it resembles the genus Zanna from
Africa and Asia. The type-species is A. peru-
ana, new species. Atiierzanna differs from
Odontoptera, which has the head similarly
shaped, by the differences in the tegmina, for
it lacks the sharp sutural angle and the spot on
the nodal line of Odontoptera.

Amerzanna peruana, n. sp.

Figs. 27, 28

Length. — Male 31-33 mm, female 35-41
mm; head 15.6 mm; ratio, head/pronotum
8.3.

Ground color reddish brown. Tegmina well
pigmented, transparent, but not obviously so
(the hind wings may be seen through the teg-
mina). Hind wings transparent, greenish
cloud at base, brown cloud at apex, brown
veins between.

HOLOTYPE (male) and ALLOTYPE (female). —
Peru: Madre de Dios: Rio Tambopata Res.,
30 air km SW Pto. Maldonada, 290 m, 16-20-
XI-1979, J. B. Heppner, subtropical, moist
forest (USNM). Paratypes (10 female, 1 male):
3, same data (USNM); 2, 11-15-XI-1979
(USNM); 2 females, 1 male 6-10-XI-1979
(USNM, LOB); 2, Loreto, Ucayali R. Yarina
Cocha, 30-XI-1954, leg. Peter Hocking, 1
head process broken (LOB, FMNH); 1,
Yahuarmayo, Ape. Group (MCZ).

Amycle Stal
Figs. 14-22, 93-101

Amtjcle Stal 1861:148. Type species: A. amahilis (West-
wood), subsequent designation by Van Duzee
1916:78

Cyrpoptus (Amyele) [sic] Stal, 1862:305.

Small, brown insects, 12-23 mm long, head
1/8 to 1/4 length of insect. Head process por-
rect, dorsoventrally flattened, dorsal surface
flat, ventral convex; usually elongate triangu-
lar, but sometimes parallel-sided, sometimes
slightly enlarged at apex; dorsal lateral carinae
sinuate, it and other carinae variable between
species. Preocular flange horizontal, about as

produced as marginal carinae. Tegmen with
M not pectinate (2 main branches), cross-
veins reticulate, apical margin more angulate
than in most genera. Metatibia with 4-5 lat-
eral spines (sometimes 3 or 7 on a single tibia).
Female ninth tergite 1.5-2 times as long as
eighth.

This genus may be separated from other
small, brown fulgorids by the porrect head
being flattened dorsoventrally. It resembles
Cyrpoptus in wing coloration. Diagnoses are
not provided; these species may be identified
by the shape of the head (Figs. 14-22) and the
color and color pattern of the hind wings
(Figs. 93-101).

Key to the Species oi Amycle

1. Hind wing with 2 colors, brown posterior and

base white, yellow, pink, or red (Fig. 93)

saxatilis Van Duzee

— Hind wing with 3 colors, brown posterior,
translucent area (Figs. 94-101), and base white,
yellow, orange, or red 2

2(1). Base of hind wing yellow, white, or pink 3

— Base of hind wing red or orange 4

3(2). Base of hind wing yellow or yellow-orange, con-
tiguous with brown suffusion, apical area lightly
suffused with brown, intermediate area translu-
cent (Fig. 101) amahilis (WesUvood)

— Base of hind wing pale pink, contiguous with
white band, posterior margin brown (Fig. 97);

sides of head process parallel (Fig. 18)

vernalis Manee

4(2). Vertex at least twice as long as broad (Figs.

17-21) 5

— Vertex less than twice as long as broad (Figs. 14,
15) 8

5(4). Hind wing with brown apical and anal areas

contiguous (Figs. 93-97) 6

— Hind wing with transparent area between
brown apical and anal areas (Figs. 98-100) 7

6(5). Hind wing with transparent area restricted to
narrow anterior band (Fig. 94); head process
parallel-sided (Fig. 20) sodalis Stal

— Hind wing with transparent area about 1/4 area
of wing (Fig. 9.5); head process subtriangular
(Fig. 21) mankinsi, n. sp.

7(5). Head process expanded at tip (Fig. 19); hind
wing with red area separated from brown anal
and apical areas by transparent band (Fig. 98)
grandis, n. sp.

— Head process subtriangular (Fig. 17); red area
and brown anal area contiguous (Fig. 100) ....
tumacacoriae Knull & Knull

8(4). Head broader than long (Fig. 14); apex of hind

wing narrowly brown (Fig. 99) brevis, n. sp.

— Head longer than broad (Fig. 15); apex of hind

142

Great Basin Naturalist Memoirs

No. 12

wing broadly brown (Fig. 96)

pinyonae Knull & Knull

Amycle amabilis (Westwood)

Figs. 22, 101

Fttlgora amabilis Westwood 1842:119. Type repository:

UMO.
Amycle amabilis (Westwood), Stal 1861: 148.

Length. — Male 14 mm; head 3.4 mm, ra-
tio, head pronotum 3. 1.

Head process elongate, parallel-sided.
Hind wing yellow to yellow-orange at base,
followed by brown suffusion, apical area less
darkly suffused, area between latter two hya-
line.

Distribution. — Holotype and paratype:
Mexico. No other specimens seen.

Amycle brevis, n. sp.

Figs. 14, 99

Length. — Male 13 mm, female 14 mm;
head 1.8 mm; ratio, head/pronotum 1.0.

Head triangular, not produced into distinct
process (Fig. 14). Hind wing pale orange at
base, anal area pale brown, dark brown area
on each side of anal fold, apical row of cells
brown (Fig. 79). Tegmina in male pale orange
with brown maculation apically; in female
mottled brown throughout, forming a pattern
with a clean, apical, diagonal line.

The two specimens upon which this species
is based may represent different species, but
the head and hind wings are similar; examina-
tion of male genitalia will be necessary to de-
termine whether these are two different spe-
cies. A. brevis may be separated from other
species by its short head.

Holotype (male). — Mexico: Nayarit, San
Bias, 4-IX-1971, W. J. Hansen (LOB). Fe-
male (not allotype or paratype): Guerrero:
Ciudad Altamirano 20-X-1983, H. Brailov-
sky, E. Barrera (LOB).

Amycle grandis, n. sp.
Figs. 19, 45, 98

Length. — Female 20-22 mm; head 5 mm;
ratio, head/pronotum 4.5.

Head elongate, slightly expanded at apex
(Fig. 19). Hind wing orange in basal 1/3,
brown in anal 1/3 and apical 1/4, translucent
areas between (Fig. 98). Tegmina brown with
most veins orange-red; two diagonal, brown
lines delineating translucent half oval on
costa, translucent triangle at apical angle, and

mottled brown and white pattern in anal angle
(Fig. 45).

Holotype (female). — Mexico: Sinaloa, Ma-
zatlan, 5-VIII-1971, D. W. Davis (LOB).
Paratype (female): Sonora, Canyon Sapopa,
15-X-1934, Rio Mayo, H. S. Gentry (CIS).
Possible male (not paratype): Jalisco, Chapala,
6 mi W Jahsco, 30-VM963, J. Doyen (LOB).

Amycle rnankinsi, n. sp.

Figs. 21, 95

Length. — Female 17 mm; head 3.5 mm;
ratio, head/pronotum 2.9.

Head slightly elongate triangular, turned
up at apex (Fig. 21). Hind wing red in basal
third, apical and anal thirds brown, all colored
areas separated from each other by pale areas
(Fig. 95). Tegmen brown with most veins red.

I dedicate this species to Dr. J. V. Mankins,
with whom we have spent many happy hours
collecting.

Holotype (female). — Honduras: Lago
Yojoa, 2-IX-1977, J. V. Mankins (LOB). Fe-
male paratype: same data except 6-X-1977
(LOB).

Amycle pinyonae Knull & Knull

Figs. 15, 96

Amycle pinyonae Knull & Knull 1947:397. Type reposi-
tory: OSU.

Length. — Male 8 mm, female 14-14.5
mm; head 2.6 mm, ratio of head/pronotum
1.9.

Head process triangular, lateral margins
sinuate. Hind wing reddish orange in basal
2/5, anal and apical angles brown, translucent
area between with brown veins. Tegmen dark
brown with many clear areas, these not form-
ing definite pattern.

Distribution. — Holotype (male): Califor-
nia: Pinyon Flat, Santa Rosa Mts. on Pinus
cembroides var. monophylla Voss. I have seen
three other specimens collected in August
from California: Pinyon Flats and Valvermo
and New Mexico: Juan Tabo, 7,000 ft.

Amycle saxatilis Van Duzee

Figs. 16, 93

Amycle saxatilis Van Duzee 1914:33. Type repository:
CAS.

Length. — Male lectotype 12.5 mm; head
2.8 mm, ratio, head/pronotum 3.5.

Head process elongate triangular. Hind
wing pale orange at base, followed by white

1988

O'BRIEN: New World Fulgoridae

143

Figs 29-45 Head lateral view: 29, Diareusa conspersa Schmidt; 30, D. kemneri Ossiannilsson; 31, D. annularis
(Olivier) 32 D imitatrix Ossiannilsson. Dorsal view: 33, Artacie haemoptera (Perty); 34, A. dufouri (Signoret). Lateral
view 35 A dufouri (Signoret) (pronotal nodules shown only in lateral view); 36, Enhydria longicornuta Lallemand; 37,
£ tesseilata (Walker); 38, E. cicadina Gerstaecker. Frontal view: 39, E. cicadina Gerstaecker Lateral view: 40,
Copidocephala merula Distant (nodules not shown); 41, C. viridiguttata Stal; 42, C. guttata white. Tegmen: 43,
Sinuala tuberculata O'Brien; 44, Odontoptera carrenoi Signoret; 45, Amycle grandis O Bnen.

144

Great Basin Naturalist Memoirs

No. 12

band, posterior half light brown. Tegmen
pale, only slightly darker at apex.

Van Duzee said the color at the base of the
hind wing varied from red to luteus in his
specimens. This has been verified (N. Penny,
personal communication). The color pattern,
however, in both specimens has no translu-
cent area.

Distribution. — Lectotype, right wing re-
moved and pinned beneath specimen: Cali-
fornia: San Diego County. I have seen no
other specimens.

Amijcle sodalis Stal

Figs. 20, 94

Amycle sodalis Stal 1861:148. Type repository: NRS.

Length. — Male 13-14 mm, female 16.5-
18 mm; head 4.5 mm, ratio, head/pronotum
4.5.

Head process elongate, sides parallel to
apex. Hind wing orange at base, dark brown
posteriorly, anterior 1 1/2 rows of cells behind
orange base translucent (Fig. 94). Tegmina
red basally, apically translucent with 2 dark
apical lines (similar to Cyrpoptiis); some dark
patches caudally.

Distribution. — Holotype: Mexico. I have
seen six other specimens collected from July
through November from El Salvador: Santa
Tecla, 900 m and Mexico: Atlixco, Canada de
Negros, Cordoba, Huachinango.

Amycle tumacacorae Knull & Knull

Figs. 17, 100

Amycle tumacacorae Knull & Knull 1947:398. Type
repository: OSU.

Length. — Male 14 mm, female 14-15.5
mm; head 3.7 mm; ratio, head/pronotum 3.7.

Head process elongate triangular, slightly
widened just before apex, slightly bent dorsad
in lateral view. Hind wing red-orange at basal
third, dark anal and apical areas, gray mem-
branous areas between with dark veins. Teg-
men dark brown with few translucent apical
cells. This species is very close to A. vernalis
in shape and length of head process; it differs
in the color and pattern in the hind wing.

Distribution. — Holotype (male): Arizona:
Tumacacori Mts. I have seen 12 other speci-
mens from Arizona: Catalina Mts., Chiric-
ahua National Monument, Molino Basin, and
Pepper Sauce Canyon; Texas: Eastland
County, Forestburg, 31 mi W Ozone, Tyler.

Amycle vernalis Manee

Figs. 18, 97

Amyele[sic] vernalis Manee 1910:117. Type repository;

USNM.

Length. — Female 14.5 mm; head 4 mm;
ratio, head/pronotum 4.0.

Head elongate, sides gradually narrowing
to apex, apex slightly turned up. Hind wing
pale pink in basal fourth, followed by band of
white, posteriorly black.

Distribution. — Lectotype: North Caro-
lina: Southern Pines (USNM). I have seen
one other specimen from Louisiana: Gal-
braith.

Aphrodisias Kirkaldy

Figs. 49, 50

Compsoptera Stal 1869:236. Type species C. cacica Stal,
original designation.

Aphrodisias Kirkaldy 1906:248, new name for Compsop-
tera Stal.

Small, reddish brown insects, 18-23 mm
long, head about 1/8 length of insect. Head
process porrect, terete, transversely striate
on all sides; dorsolateral carinae present;
short, nonstriate, apical portion with dorsal
median carina and longitudinal ridge on verti-
cal face. Preocular flange horizontal. Tegmina
with longitudinal veins irregularly angled, not
straight; few cross-veins. Female ninth tergite
slightly longer than eighth.

This genus may be separated from all other
genera by the wrinkled, porrect head process.
The head process of Artacie also is textured,
but it is recurved and recumbent on the ver-
tex.

Key to the Species of Aphrodisias

1. Head with wrinkled or striate portion of process
shorter than pronotum; hind wings red at base,
brown apically cacica (Stal)

— Head with wrinkled portion of process longer
than pronotum; hind wing red, white, and brown
shaman, n. sp.

Aphrodisias cacica (Stal)
Fig. 49

Compsoptera cacica Stal 1869:237. Type repository:

NRS.
Aphrodisias cacica (Stal), Kirkaldy 1906:248.

Length. — Male 18-20 mm; head 2 mm;
ratio, head/pronotum 1.5.

Head process siibe(|ual in length to vertex.
Hind wing red at base, brown posteriorly.
Tegmen brown.

1988

OBrien; New World Fulc;ohidae

145

Figs. 46-62. Head and pronotum, dorsal view: 46, Sinuala stali O'Brien; 47, S. schmidti O'Brien; 48, S. tuberculata
O'Brien; 49, Aphrodisias cacica (Stal); 50, A. shaman O'Brien. Head and thorax, dorsal view and lateral view, and head,
ventral view: 51-53, Chilobia silena Stal; 54-56, C. smaragadina (Walker); 57-59, C. cinxia Stal; 60-62, C.
dichopteroides (Distant).

146

Great Basin Naturalist Memoirs

No. 12

Distribution. — Holotype (female): Mex-
ico. I have seen six specimens collected from
Mexico: Alamos, 15 mi S Cuernavaca, Cuer-
nonica, from September through November.

Aphrodisias shaman, n. sp.
Fig. 50

Length. — Female 23 mm; head 3.5 mm;
ratio, head/pronotum 2.3.

Head process 1.7 x longer than vertex.
Hind wing pale pink at base, triangular, white
area intruding into pink, apex and anal area
missing, but edges of anal area brown, apical
area brown with white spots.

The wings of this specimen are badly dam-
aged, but it may be separated from cacica on
the basis of head shape. This specimen has lost
its left tegmen and hind wing and part of the
right wing, but it is a very distinctive species
based on the head alone.

Holotype (female). — Mexico: Sinaloa, 5
mi N Mazatlan, l-VII-1965, J. A. and M. A.
Chemsakand E. G. and J. M. Linsley (UCB).

Artacie Stal
Figs. 33-35

Artacie Stal 1866:132. Type species: Flata haemoptera
Perty, subsequent designation by da Costa Lima
1935:488.

Brown-winged, medium-sized insects, 20-
27 mm long. Head process terete, recurved to
lie against vertex (Fig. 35), texture of apex
unevenly, rugosely tuberculate, ventral me-
dian and marginal carina on process, lateral
carinae on frons. Pre- and supraocular flanges
large (Figs. 33, 34). Pronotum with white
nodules. Tegmina with main veins slightly
raised, surface concave between them, M 3-
branched, cross- veins reticulate in corium,
forming rectangular cells in membrane. Red
or orange-red color at base of hind wing al-
most reaching apical margin medially, divid-
ing it into apical and anal areas.

I synonymized the two species in this genus
in 1985. After studying additional specimens,
I conclude that I was in error and that the
artist's illustrations of the species were correct
in each case and there is a difference in the
shape of the vertex of the two species. The
white spots in the apical portion of the wings
do vary in size, placement, and number be-
tween specimens, but a general pattern for
haemoptera is described below.

Key to the Species oi Artacie

1. Median length of head process before preocular
flange subequal to median length of vertex ....
dufourii (Signoret)

— Median length of head process before preocular

flange half of median length of vertex

haemoptera (Perty)

Artacie dufourii (Signoret), new status
Figs. ,34, 35

Encophora [sic] dufourii Signoret 1858:497, pi. 12, no. 2,

Fig. 1. Type repository: MZF?
= Artacie hemoptera (Perty), O'Brien 1985:661. Error.

Length. — Male 25 mm, female 27 mm;
head length 2.6 mm; ratio, head/pronotum
1.4. Length of head process in front of preocu-
lar flange subequal to length of vertex.

Wings orangish red at base, white spots
scattered through brown apical and anal ar-
eas, not forming a figure with an empty center
(a square, circle, or diamond).

The two known species may be separated
by the proportion of the head process in front
of the preocular flange to the vertex, subequal
in dufourii and about half in haemoptera
(Figs. 33, 34).

Distribution. — Type (not seen): Cayenne.
I have seen two other specimens collected in
March from French Guiana: Maron River;
Guyana: Wineperu. Part of Signoret's collec-
tion went to the Naturhistorisches Museum,
Vienna, and part to Museo Zoologico
dell'Universita degli Studi di Firenze, Flo-
rence, according to Horn and Kahle (1935). I
did not see the type in Vienna and have not
heard from Florence.

Artacie haemoptera (Perty)
Fig. 33

Flata haemoptera Perty 1833:176, pi. 35, Fig. 3. Type

repository: ZSBS, not found.
Artacie haemoptera (Perty), Stal 1866:389.
Encophora [sic] dufourii Signoret, synonymized O'Brien

1985:661. Error.

Length.— Male 20-22 mm, female 23-25
mm; head length 2.1 mm; ratio, head/prono-
tum 1.2.

Length of head process in front of preocular
flange half length of vertex. Wings red at base,
brown in apical and anal areas, 4-6 white
spots in circle, diamond, or rectangle on api-
cal area, few smaller dots also.

These two species may be separated by the
length of the head process and the pattern of
white dots on the hind wing.

1988

O'BRIEN: New World Fulgoridae

147

Distribution. — Holotype (not seen) : Ama-
zon basin [Brazil?]. I have seen nine other
specimens collected in March, August, and
October from Brazil: Canum, Manaus,
Obidos, Rio Negro, and Teffe. The type could
not be found in the Zoologische Sammlungen
des Bayerischen Staates, Munich, in 1975.

Cathedra Kirkaldy

Fig. 26

Cathedra Kirkaldy 1903:179. Type-species: Phrictus ser-
rata (F.), original designation.

Pristiopsis Schmidt 1905:332. Type-species: Fulgora ser-
rata F. , original description; synonymized by Dis-
tant 1906:20.

Large, yellowish brown insects, 52-71
mm, head 1/3 length of insect. Head process
porrect, terete, gradually narrowed to apex;
marginal and lateral carinae of frons produced
into 11 and 8 spines, respectively; head about
1/3 length of insect. Preocular flange horizon-
tal, spinelike. Tegmen with M irregularly
branched (2 main branches); clavus open;
cross-veins forming almost round cells anteri-
orly, arched and parallel behind nodal line.
Hind wings brown with ochre eye spot at
apex, pale, elongate spots at base, some waxy
spots and dashes throughout; emarginate on
posterior border before spot.

This monotypic genus has a head process
like a cathedral spire, with many pairs of lat-
eral spines, which distinguishes it immedi-
ately from any other fulgorid genera.

Cathedra serrata (Fabricius)

Fig. 26

Fulgora serrata Fabricius 1781:313.

Phrictus serratus (Fabricius), Schaum 1850:65.

Cathedra serrata (Fabricius), Distant 1906:20.

Length. — Male 52-53 mm, female 61-71
mm; head 19 mm; ratio head/pronotum 6.3.

Ground color ochre mottled with brown.

Distribution. — Holotype: Suriname. I
have studied eight other specimens collected
from January to November from Bolivia:
Santa Cruz; Brazil: Manaus-Itacoatiana road,
km 30; Panama: Barro Colorado Island. Zim-
sen (1964) did not list the repository of this
Fabrician type.

Chilobia Stal
Figs. 51-62

Chilobia Stal 1863:237. Type-species: C. cinxia Stal, sub-
sequent designation by da Costa Lima 1935:497.

Ecuadoria Distant 1906:21. Type-species: E. dichop-
teroides Distant, original designation. New syn-
onymy.

Tegmina membranous throughout or mem-
branous in apical half, basal half opaque,
brown or brownish red, medium-sized in-
sects, 19-28 mm long. Head process usually
recurved, usually dorsoventrally compressed,
usually gradually expanding in ventral view
towards apex. Lateral carinae of frons parallel
or diverging to near base, carinate throughout
or becoming broad, fused with median carina
into broad hump. Frons elevated above level
of frontoclypeal suture, expanded into lobes
near apex, with transverse carina extending
across frons from lobe to lobe. Vertex with
lateral carinae sinuate or angulate in lateral
view, supraocular flanges present, apical mar-
gin V-shaped, sometimes angulately emar-
ginate medially in addition, sometimes hid-
den under recurved process. Preocular flange
absent. Pronotum with few white nodules.
Tegmen with M pectinate (5-8 branches);
clavus closed; cross-veins irregular and dense
in basal half, few and as thick as longitudinal
veins in apical half; longitudinal veins with
small spines alternately angled to left or right;
transition zone between membrane and
opaque area, when present, is arched. Hind
wings variable, from only slightly enfumed at
base to bright red with sinuate, brown, con-
tiguous area, only two marginal rows of cells
enfumed at apex. Female ninth abdominal
tergite shorter than eighth.

Because I have intermediates and because
the male genitalia are very similar, I have
synonymized t\i^ genera Ecuadoria and En-
hydria in spite of several character differ-
ences. The species of Ecuadoria have a long
head process and brightly colored tegmen and
hind wing, whereas the Enhydria species
have a short head process, transparent teg-
mina, and clear hind wings.

The four described species are illustrat-
ed, but 1 have not provided a key because I
am unable to determine the range of varia-
tion within a species due to lack of available
material.

Chilobia dichopteroides (Distant), n. comb.
Figs. 60-62

Ecuadoria dichopteroides Distant 1906:22. Type reposi-
tory: BMNH.

148

Great Basin Naturalist Memoirs

No. 12

Distribution, — Holotype: Ecuador (nw):
Rio Durango, 350 ft. I have seen no other
specimens.

Chilobia cinxia Stal

Figs. 57-59

Chilobia cinxia Stal 1863:238. Type repository: NHMV.

Distribution. — Holotype (female): Vene-
zuela. I have seen no other specimens.

Chilobia silena Stal
Figs. 51-53

Chilobia silena Stal 1863:238. Type repository: NRS.

Distribution. — Holotype (male): Ecua-
dor: Quito. I have seen no other specimens.

Chilobia smaragdina (Walker)
Figs. 54-56

Dichoptera smaragdina Walker 1851:304. Type reposi-
tory: BMNH.
Chilobia smaragdina (Walker), Gerstaecker 1895:39.

Distribution. — Holotype: Venezuela. I
have seen no other specimens.

Copidocephala Stal
Figs. 40-42

Copidocephala Stal 1869:235. Type-species: Enchophora
guttata White, original designation.

Coanaco Distant 1887:28. Type-species: Enchophora
guttata White, original designation; synonymized
by Distant 1906:23.

Medium-sized, greenish or brown or red-
dish brown insects, 22-32 mm long. Head
process recurved at junction with head, later-
ally compressed, gradually narrowing ante-
rad; lateral carinae of frons fused at junction
with process with medium and lateral carinae
of process into one large mound; dorsolateral
carina also present on process. Preocular
flange absent. Pronotum with small, white,
hemispherical tubercles, median carina not
strongly humped. Tegmen with M usually
pectinate (5-6 branches); corium with cross-
veins regularly, evenly reticulate, with
smooth, sunken, red spots without cross-
veins; membrane with cells subsquare. Fe-
male eighth and ninth abdominal tergites
subequal in length.

This genus has the head process curved
back at the base, never extending foi-ward as
in some Enchophora; it is scimitar-shaped like
Enchophora and Enhydria. Copidocephala
differs from Enhydria, which has membran-

ous tegmina, and from Enchophora in the
characters of the thorax given in that generic
discussion. The close proximity of the prono-
tum and eyes suggests that the postocular
flange serves to protect the eye from contact
with the pronotum.

Key to the Species of Copidocephala
1. Hind wing black, luispotted merula Distant

— Hind wing brown, spotted with red or bine,
green or white 2

2(1). Large species, 28 mm or longer; hind wing

brown with red spots guttata (White)

— Small species, 25 mm or less; hind wing brown

with blue, green, or white spots

viridiguttata Stal

Copidocephala guttata (White)
Fig. 42.

Enchophora guttata White 1846:331. Type repository:

BMNH.
Copidocephala guttata (White), Stal 1869:236.

Length. — Female 29-32 mm, head length
1.4 mm; ratio, head/pronotum .63. Pronotum
and costal margin of tegmen green, pronotum
with red, transverse, sometimes broken,
band; ground color of rest of insect ranging
from yellow or green through red to olive
green. Corium with approximately 21 round,
smooth, sunken, small (1 mm or smaller), red
spots; spots differing slightly in number, size,
and placement among individuals. Hind wing
brown with red spots in basal two-thirds.

This species may be separated from inerida,
which is similar in size and coloration, by the
absence of red spots in the hind wing in
merula. It differs from viridiguttata in size
and in the color of spots in the hind wing, blue
or green to white spots in viridiguttata.

Distribution. — Type: South America. I
have seen 20 other female specimens col-
lected from March through September,
November and January. Panama: Barro Colo-
rado Island, Chiriqui Provence, Fort Clayton
Costa Rica: 6 mi W Las Canas, La Selva
Honduras: Catacamas, 15 km W La Ceiba
Mexico: Tamazunchale, Temiscal, Volcan
Tacana.

Copidocephala merula Distant

Fig. 40

Copidoccpluda iiicrula Distant 1906:23. Type repository:
BMNH.

Coanaco melanoptera Schmidt 1907:361. T\pe reposi-
tory: IZW.

1988

O'BRIEN; New World Fulgoridae

149

Copidocephala melanoptera (Schmiclt), Metcall 1947:
176, New synonymy.

Length. — Female, 34-38 mm.

Ground color as in guttata except hind wing
black, without red spots.

The differences in coloration of the tegmen
found in the two specimens of mcrula and
melanoptera are within the variation found in
the species guttata; so on the basis of size and
the color of the hind wing, I synonymize the
two species.

Distribution. — Holotypes: Colombia (both
types). I have seen the type oimerula but not
o{ melanoptera. No other specimens seen.

Copidocephala viridiguttata Stal
Fig. 41

Copidocephala viridiguttata Stal 1869:236. Type reposi-
tory: NHMV.

Coanaco ornanda Di.stant 1887:29, pi. 4, Fig. 13. Type
repository: BMNH.

Copidocephala ornanda (Distant), da Costa Lima 1935:
491. New synonymy.

Length. — Male 22-24 mm; head 1.1 mm;
ratio of head/pronotum 0.5.

Ground color olive green with black or red-
black, sunken, smooth, round spots in same
pattern as guttata. Hind wing brown with
blue or green to white spots.

This species differs from the other two in
having blue, green, or white spots in the hind
wing rather than red spots or none. It is inter-
esting that I have 13 males of this species for
study and 20 females of guttata. I will examine
other collections to see if this is coincidence or
unexpected sexual dimorphism. I synonymize
ornanda on the basis of the shape of the head
process and wing coloration.

Distribution. — Type: Golombia (viridi-
guttata); syntypes: Panama: Bugaba, Tole (or-
nanda). I have studied 13 other specimens
collected from May through September and
January from Panama: Achiote Road, Barro
Colorado Island, Fort Clayton; Costa Riga:
Golfito, La Selva; Mexico: Los Tuxtlas, 3 mi S
Palenque.

Diareusa Walker

Figs. 29-32

Diareusa Walker 1858:43. Type-species: Phrictus annu-
laris (Olivier), by monotypy.

Large, reddish brown insects, 32-49 mm,
head relatively short, from .1 to .2 length
of the insect. Head process porrect, terete,

gradually slightly enlarging at apex; median
ventral carina in about anterior third of pro-
cess, ventral and dorsal lateral carinae com-
plete; short dorsal median carina in anterior
portion. Preocular flange vertical, well pro-
duced. Tegmen with M sometimes pectinate
(3-4 main branches), sometimes evenly
branched; cross-veins irregular in corium,
forming small squares beyond stigmal line.
Tegmen brown throughout with some ante-
rior cells and many spots either white or red;
often some apical spots translucent. Female
eighth and ninth abdominal tergites subequal
in length.

Diareusa are large, reddish brown insects
with a comparatively short (.1 to .2 length of
insect), terete, but not inflated head without
transverse striae or spines; these characters
will separate them from other New World
Fulgoridae. The species differ in the color of
spots in the hind wing and in the shape of the
head process.

Key to the Species of Diareusa

1. Spots on hind wing red and white

conspersa Schmidt

— Spots on hind wing white only 2

2(1). Head process short; vertex and head process as

long as width of head, including eye

kemneri Ossiannilsson

— Head process longer than width of head includ-
ing eye (Fig. 30) 3

3(2). Dorsal surface of head process in lateral view

convex (Fig. 31) annularis (Olivier)

— Dorsal surface of head process in lateral view

straight or concave, not convex (Fig. 32)

imitatrix Ossiannilsson

Diareusa annularis (Olivier)

Fig. 31

Fulgora annularis Olivier 1791:568. Type repository: un-
known.
Phrictus annularis (Olivier), Walker 1851:264.
Diareusa annidaris (Olivier), Walker 1858:44.

Length. — Male 32-38 mm, female 45 mm;
head 6.2 mm; ratio of head/pronotum 2.2.

Head process in lateral view convex dor-
sally (Fig. 31.) Hind wings with white spots.
These two characters separate it from the
other species.

Distribution. — Type (not seen): Suri-
NAME. I have seen 10 other specimens col-
lected June to August and January from PERU:
Brazilian frontier; BRAZIL: Serra do Navio,
Sinop; French Guiana: 67 km S Cayenne;
SURINAME: Mapane; GUYANA: Shudihat River.

150

Great Basin Naturalist Memoirs

No. 12

Diareusa conspersa Schmidt

Diareusa conspersa Schmidt 1906:375. Type repository:

IZW.
Diareusa dahli Ossiannilsson 1940:44, Fig. 2. Type

repository: UZIL. Tslew synonymy.

Length. — Male 38 mm, female 46-49 mm;
head 5.3 mm, ratio of head/pronotum 1.8.

Hind wings with red spots. Head process
similar to that odinitatrix.

This is the most easily identified species in
the genus with the spots on the hind wing red
and white; in all other species they are white.
It is interesting that although this is the easiest
character for identification, both Schmidt and
Ossiannilsson described this character among
the last, causing it to be overlooked. Thus,
many museums have imitatrix, the common
Central American species, identified as con-
spersa. I synonymize conspersa and dahli on
the basis of head shape and wing color.

Distribution. — Holotypes: Ecuador: Pal-
mar (conspersa); Colombia: Rio San Agustin
(dahli). I have seen three other specimens
collected from June through August from
Ecuador: Amhuagu, Rio Palenque.

Diareusa imitatrix Ossiannilsson

Fig. 32

Diareusa itnitatrix Ossiannilsson 1940:45, Fig. 3. Type
repository: UZIL.

Length. — Male 35-39 mm, female 43-48;
head 5.8 mm; ratio head/pronotum 2.2.

Hind wing with white spots. Head process
similar to conspersa, but longer and narrower
than other species with white-spotted hind
wings.

This is the only Central American species
and often has been misidentified as con-
spersa. Panamanians call this insect "totoran.

Distribution. — Holotype (female): Pan-
ama: Chiriqui. I have seen 50 other specimens
collected all year around from Colombia:
Turbo; Venezuela: Rancho Grande; Panama:
Barro Colorado Island, Fortuna, Hartman's
Finca, Margarita; Costa Rica: La Selva, Tur-
rialba; Belize: Middlesex; Mexico: Bonampa
R., 1 mi W Fortin de las Flores, Los Tuxtlas.

Diareusa kemneri Ossiannilsson

Fig. 30

Diareusa kemneri Ossiannilsson 1940:43, Fig. 1. Type
repository: UZIL.

Length. — Male 35 mm; head 4 mm; ratio.

head/pronotum 1.8.

Hind wings with white spots. Head process
short (Fig. 30). D. kemneri may be separated
from other white-spotted species by the short
head process.

Distribution. — Holotype (female): Peru:
Perene. I have studied one other specimen
collected in September from PERU: Aguaitia.

Enchophora Spinola

Figs. 63-76

Enchophora Spinola 1839:221. Type-species: recurva,
subsequent designation by Duponchel 1840:200.

Medium-sized, reddish brown, green, yel-
low, or mottled green, white, and red insects,
22-32 mm long. Head process recurved, lat-
erally compressed; nine carinae present on
process (all except dorsal median carina). Pre-
ocular flange present as vertical carina linking
frons and vertex laterally (Fig. 3). Pronotum
with pair of deep fossettes laterad of strongly
raised median carina. Apex of mesonotum
with striate depression between bifurcate me-
dian carina. Tegmen with few strong, longitu-
dinal veins, cross-veins variable, dense to av-
erage. Female abdominal segments 8 and 9
subequal in length.

Enchophora may be separated from the
very similar Copidocephala by the raised me-
dian carina of the pronotum with a fossette or
depression on each side, the depression at the
apex of the mesonotum surrounded by a high
ridge, and the ventrolateral fields of the
pronotum being separated from the eye by a
space about the width of the last antennal
segment. In Copidocephala the callus behind
the eye almost touches the pronotum when
viewed from above, and the pronotum and
mesonotum are smoothly rounded. Both gen-
era lack the preocular flange. Enhijdria,
which also has a scimitar-shaped head pro-
cess, may be differentiated by its pale tegmen
and small spines on the tegminal veins.

The species may be separated by the shape
of the head process (although this structure
varies slightly in curvature and compression),
the color of the hind wings, and sometimes by
distinctive marks or colors on the body or
wings. Waxy spots on the wings may be help-
ful if they have not been rubbed off; often they
may be identified by a smooth, pale brown
area if the wax is missing. The hind wings are
usually red, yellow, orange, or white in the
basal 1/4 to 2/3, measured along the anterior

1988

OBrien: New World Fulgoridae

151

margin, or 5/6 if measured from anterior to
posterior. The color pattern of the tegmen
may be quite variable, as in sanguinea Dis-
tant, nigromaculata Distant, and reciirva
(Olivier) where, in females, the tegmen may
be concolorous, marked with discrete red or
black spots, or marked with coalescent red
spots. Although I have not seen both sexes of
every species, if there is a sexual difference in
color pattern, the male tends to be more
evenly mottled and less variable.

Three species listed as Enchophora in Met-
calf's (1947) catalog have been transferred
since to other genera (see list of species).

Key to the Species of Enchophora

1. Hind wing with white, waxy points on red

area stillifera (Stal)

— Hind wing without white, waxy points on red
area 2

2(1). Head process about twice length of frons be-
fore process, enlarged and narrowed near tip
(Figs. 63, 64) .3

— Head process less than twice length of frons
before process, scimitar-shaped, curved back
over vertex, or curved and then bent upward
4

3(2). Apex of head process black, broadly trilobed

(Fig. 63) reciirva (Olivier)

— Head process brown throughout, narrowly
expanded at apex (Fig. 64) tuba (Germar)

4(2). Hind wing red in basal half; posterior half
medium to dark brown; tegmen brown or
reddish brown, often mottled 5

— Hind wing white, pale orange, or brown at
base; either only apical 1/4 pale brown or
posterior half medium brown (subviridis, tu-
berculata), tegmen unmottled green or yel-
low or green mottled with red 9

Apex of pronotum with transverse, black
band (Fig. 66) nigromaculata Distant

Apex of pronotum without transverse, black
band 6

Mesonotum with black spots (Figs. 67, 76) ... 7

Mesonotum without black spots 8

Mesonotum with pair of black spots, also pair
on ventrolateral fields of pronotum; head pro-
cess recurved to near vertex, then bent up-
ward (Fig. 67) pyrrhocrypta Walker

— Mesonotum with 2 or 3 pairs of black spots;
head process short, only slightly recurved
(Fig. 76) maculata, n. sp.

8(6). Apical fourth of pronotum pale; base of mem-
brane of tegmen with a brown spot about size
of eye on each side, 3-5 other such spots
usually present on membrane; ca basal half of
hind wing red; abdominal tergites brown
pallidiptinctata Lallemand

5(4).

6(5).

7(6).

— Pronotum concolorous; tegmen not as above;
ca basal 2/3 of hind wing red; abdominal ter-
gites bright red sanguinea Distant

9(4). Hind wing white at base 10

— Hind wing yellow, orange, or brown at base
12

10(9). Tegmen green or yellowish 11

— Tegmen greenish white mottled with red . .
rosacea Distant

11(10). Tegmen green or yellowish with white costal

margin; head process large (Fig. 71)

prasina Gerstaecker

— Tegmen greenish yellow throughout; head
process small (F'ig. 72) uniformis, n. sp.

12(9). Head process scimitar-shaped (Fig. 74); hind
wing yellow at base, then white, then brown
at apex viridipennis Spinola

— Head process recurved and wrinkled (Figs.
73, 75); hind wing orange at base, brown at
apex 13

13(12). Head process recurved, then bent upward at

apex (Fig. 75) subviridis Distant

— Head process recurved almost to vertex (Fig.
73) tuberculata (Olivier)

Enchophora maculata, n. sp.

Fig. 76

Length. — Male 24 mm, female 28 mm.

Ground color brown with minute dots of
red and green. Head process short, extending
forward then dorsad (Fig. 76). Wings brown
on apical third, red basally, red extending
from anterior to posterior margin. Distinctive
markings: two or three pairs of brown spots on
pronotum (Fig. 76); anterior spot black, poste-
rior medium brown, sometimes pale brown
spot laterad of latter. Abdominal tergites
brownish red.

This species may be identified easily by its
unusual, short, partially forward-directed
head process (Fig. 76) and the spots on the
pronotum.

HoLOTYPE (male) and allotype (female). —
Peru: Madre de Dios: Rio Tambopata Res.,
30 air km SW Pto. Maldonada, 290 m, 16-20-
Xl-1979, J. B. Heppner, subtropical, moist
forest (USNM).

Enchophora nigromaculata Distant

Fig. 66

Enchophora nigromaculata Distant 1906:23. Type repos-
itory: BMNH.

£. nigrolimbata Lallemand 1938:350. Type repository:
DEI. New synonymy.

Length. — Female 24-26mm.

152

Great Basin Naturalist Memoirs

No. 12

Figs. 63-76. Head and pronotum, lateral view, ofEnchophora: 63, E. recurva (Olivier); 64, E. tuba (Germar); 65, £.
stillifera (Stal); 66, E. nigromaculata Distant; 67, E. pyrrhocrypta Walker; 68, £. piillidipunctata Lallemand; 69, E.
sanguinea Distant; 70, E. rosacea Distant; 71, E. prasina Gerstaecker; 72, £. uiiiformis O Brien; 73, E. tuhercidata
(Olivier); 74, E. viridipennis Spinola; 75, E. subviridis Distant; 76, E. maculata O'Brien.

Ground color reddish brown, variable, teg-
men may be clear, mottled with yellow veins,
or with 10 dark spots; membrane brown or
with dark spots, 5 to 7 waxy points along nodal
line. Head process scimitar-shaped (Fig. 66).
Wings red at base, brown posteriorly.

This species may be separated from all
other Enchophora by the black line along the
hind margin of the pronotum. I have not seen
the type o(nifi,rolimhata.

Distribution. — Holotype: Bolivia (nigro-
maculata), Peru (nigrolimbata). I have seen

1988

O'BRIEN: New World Fulgoridae

153

11 other specimens collected from October to
January from Bolivia: Chapare; Brazil: Rio
de Janeiro; Peru: Chanchamayo; 30 km SW
Puerto Maldonado, 290 m; Satipo; Tingo
Maria; Explorama Lodge; Ecuador: Parque
Nacional St. Cecelia.

Enchophora pallidipunctato Lallemand

Fig. 68

Enchophora pallidipunctata Lallemand 1966:.52. Type
repository; FSAG.

Length. — Male 23 mm, female 24-26 mm.

Ground color reddish brown, mottled.
Head process scimitar-shaped (Fig. 68).
Wings red on basal half, brown posterior half
(not the usual 2/3 red, 1/3 brown). Seventeen
waxy tufts present in apical area. Distinctive
markings: pale band along apical fourth of
pronotum and five round, dark spots in tegmi-
nal membrane, one on each side just behind
nodal line, three to five distad of these. Ab-
dominal tergites brown.

This species may be separated from all
other Enchophora by the pale apical fourth of
the pronotum, the dark spots in the membra-
nous area of each tegmen, and the smaller,
red portion of the hind wings. It most closely
resembles sanguinea, which has the abdomi-
nal tergites bright red, while they are brown
in pallidipunctata.

Distribution. — Holotype: Bolivia: Buena
Vista. I have seen 24 other specimens col-
lected from February through April, August
and September from Bolivia: Buena Vista,
Santa Cruz; Peru: Chanchamayo, Chontilla,
Iquitos, Lima, Pileopota, 600 m, Rio San-
tiago, Satipo, Tarapoto. Several specimens la-
beled on Copaifera Dourajeanni (Legumi-
nosae) at Tingo Maria, but I am unable to find
this species listed in any botanical reference.

Enchophora prasina Gerstaecker

Fig. 71

Enchophora prasina Gerstaecker 1895:37. Type reposi-
tory: emau.

Length. — Male 24-27 mm, female, 28
mm.

Ground color green, with costal margin of
tegmen white and head and genitalia reddish
brown. Head process scimitar-shaped, long,
almost as long as mesonotum and pronotum
combined. (Fig. 71). Wings white. No waxy
tufts evident.

This species may be separated from all
other Enchophora by the green tegmen with
white costal margin.

Distribution. — Holotype: Colombia: Nova
Granada. I have seen 12 other specimens col-
lected from April through June, and October
from Bolivia: Buena Vista; Peru: NE;
Panama: Barro Colorado Island, Cerro Azul,
Santa Rita Ridge; Costa Riga: La Selva.

Enchophora pyrrhocrypta Walker

Fig. 67

Enchophora pyrrhocrijpta Walker 1851:272. Type repos-
itory: BMNH.

Length. — Male 19-20 mm, female 23-25
mm.

Ground color reddish brown, mottled.
Head process recurved over pronotum, then
bent upward (Fig. 67). Wings red at base,
brown posteriorly. Two pairs of distinctive,
black spots, one on each side of mesonotum,
and one on each ventrolateral margin of
pronotum (Fig. 67). No waxy tufts evident.

This species may be separated from all
other Enchophora by the black marks on the
lateral fields of the pronotum and on the sides
of the mesonotum (Fig. 67). E. subviridis hsis
a similarly recurved head process.

Distribution. — Holotype: Brazil: Para. I
have seen seven other specimens collected
from October through February from Brazil:
Benjamin Constant, Rio Negro, Belem, Man-
gabeira; Guyana: Kartabo, St. Laurent;
Venezuela: Cerro de Neblina.

Enchophora recurva (Olivier)
Fig. 63

Fulgora recurva Olivier 1791:34. Type repository: un-
known.

Enchophora recurva (Olivier), Spinola 1839: 222, pi. 10,
Figs. 1, 2

Enchophora bohemani Stal 1854:244. Type repository:
NRS. New synonymy.

Length. — Male 24-26 mm.

Ground color reddish brown, tegmen mot-
tled, often with spots composed of yellow
clumped veinlets; membrane with long yel-
low and brown markings. Head process elon-
gate, produced in club that at apex is twice
average width of process, apex trilobed, black
(Fig. 63). Wings red at base, brown posteri-
orly. No waxy tufts evident.

This species may be separated from all
other Enchophora by the enlarged, blackapex
of the head process.

154

Great Basin Naturalist Memoirs

No. 12

Distribution. — Holotypes: Brazil {bohe-
manni) and Suriname (recurva). I have seen
nine other specimens from BRAZIL: Linhares,
Araxa, and Painaivas, taken from October
through December. I am unable to confirm
Berg's report from Argentina (Berg 1879).

Enchophora rosacea Distant

Fig. 70

Enchophora rosacea Distant 1887:27, pi. 4, Fig. IL Type
repository: BMNH.

Length. — Male 22-23 mm, female 25-26
mm.

Ground color green, tegmen at base green-
ish white with distinctive clumps of red dots
between the green veins; membrane green
with 20 white, waxy spots. Head process scim-
itar-shaped (Fig. 70). Wings mostly white,
dusted with red spots at base, brown posteri-
orly.

This species may be separated from all
other Enchophora by the red and green com-
bination on the tegmen.

Distribution. — Holotype: Nicaragua:
Chontales. I have seen 10 other specimens
collected from March through August from
Panama: Barro Colorado Island; Costa Riga:
La Selva.

Enchophora sangtiinea Distant
Fig. 69

Enchophora sanguinea Distant 1887:27, pi. 4, Fig. 16.
Type repository; BMNH.

Enchophora florens Distant 1887:28, pi 4, Fig. 12. Type
repository: BMNH. New synonymy.

Enchophora longirostris Distant 1887:28. Type reposi-
tory: BMNH. New synonymy.

Length.— Male 22-23, female 24-25 mm.

Ground color greenish to reddish, tegmen
greenish or reddish, or greenish black with
orange or red spots encircled with yellow
veins, or the red spots may be fused into large
areas or confluences, usually very mottled
with clumps of yellow veins and dark and/or
light spots. Head process scimitar-shaped
(Fig. 69). Wings red at base, brown posteri-
orly. Twenty waxy tufts present in membra-
nous area of tegmen. Abdominal tergites
bright red.

I synonymize this species with two other
species; E. florens and longirostris have the
same male genitalia; E. sanguinea differs from
them only in the pattern of the tegmen. When
I found comparable difference in coloration in

other species, I felt confident in synonymizing
the three. This species may be separated from
all other Enchophora with red hind wings by
the shape of the head and the lack of distinc-
tive identifying marks such as bands and spots
on the pronotum.

Distribution. — Type: Guatemala,
Panama (sanguinea). Nicaragua: Chontales
(florens); Costarica: Cache. Colombia (lon-
girostris). I have seen 39 other specimens col-
lected from July to September, with one in
February. Ecuador: Palmar; Panama: Barro
Colorado Island, Cana 1,200 ft; Costa Riga: 1
mi N Tucurrique, Rincon, Turrialba, Cairo.

Enchophora stillifera (Stal)
Fig. 65

Phrictus stillifera Stal 1862:303. Type repository:

nhmv.

Enchophora stillifera (Stal), Stal 1864:49.

Length. — Male 25-28 mm, female 32 mm.

Ground color greenish brown, tegmen of-
ten with spots of clumped yellow veins, also
often with broad patches of waxy exudate.
Head process elongate, scimitar-shaped,
about twice as long as frons below process.
Wings red at base, mottled with white, waxy
spots; brown posteriorly.

This distinctive species is the only En-
chophora with many white, waxy spots on the
red portion of the hind wing.

Distribution. — Holotype (male): Mexico;
Guatemala: Sabo; Panama: Bugaba, 2,000-
3,000 ft. I have seen 24 other specimens col-
lected from June to October from Panama
Chiriqui, Lino; CoSTA RiCA: Osa Peninsula
Honduras: Orina del Rio Camgre, Juticalpa
Belize: Northern Road, mi 28-45; Mexico
San Quintin.

Enchophora subviridis Distant

Fig. 75

Enchophora subviridis Distant 1887:28, pi. 4, Fig. 17.

Type repository: BMNH.
Enchophora subviridis vdT. a. Distant 1887:497.
Enchophora subviridis distanti Metcalf 1938:358.

Length. — Male 22 mm, female 23-25 mm.

Ground color green, brown, or yellow.
Head process reflexed, then turned up at tip.
Wings orange in basal half, brown posteriorly;
8 waxy tufts in costal margin of tegmen, 20
plus in membrane.

This species may be separated from all
other species but pijrrhocnjpta by the re-
curved and bent dorsad head process, and

1988

O'BRIEN; New World Fulcoridae

155

pyrrhocrypta has distinctive, black spots on
the pro- and mesonota and red on the hind
wings.

Distribution. — Syntypes: Panama: Chiri-
qui, Bugaba. I have seen six other specimens
collected from August to October fiom
Panama: Chiriqui, Bugaba, 800-1,500 ft;
Costa Rica: Puntarenas Prov., Osa Penin-
sula, Rincon; San Vito, Golfito.

Enchophora tuba (Germar)

Fig. 64

Fulgora tuba Germar 1830:46. Type repository: Lvov.

(All types there are lost.)
Enchophora tuba (Germar), Burmeister 1845:4.

Length. — Female 30 mm.

Ground color reddish brown, mottled.
Head process elongate, produced in club that
at apex is twice average width of process, apex
trilobed, brown like rest of process (Fig. 64).
Wings red at base, brown posteriorly. No
waxy tufts evident.

This species may be separated from all
other Enchophora by the long, thin head pro-
cess with a trilobed, concolorous tip.

Distribution. — Type: Brazil. I have seen
two other specimens collected from Brazil:
Agua Preta, in December.

Enchophora tuherculata (Olivier)

Fig. 73

Fulgora tuherculata Olivier 1791: 569. Type repository:
unknown.

Enchophora tuberculata (Olivier), Burmeister 1845: [4].
Enchophora tuberculata sub. sp. fuscomaculata
Lallemand 1956:6. Type repository: FSAG.

Enchophora parvipennis Walker 1858:30. Type reposi-
tory: BMNH. New stjnonyinij.

Length. — Male 22 mm.

Ground color greenish brown. Head pro-
cess recurved toward vertex (Fig. 73). Wings
brown, orange in anterior four cells. No waxy
tufts evident.

This species may be separated from all
other Enchophora by the recurved head pro-
cess, which differs from pyrrhocrypta and
subviridis by not being turned up at the tip.
The basal colored area of the hind wing is also
more reduced than in any other species; in
one specimen it is brown with the four cells
slightly paler brown.

Distribution. — Type: Suriname {tuhercu-
lata), Brazil: Para {parvipennis male). I have
seen two other specimens from French

Guiana: St. Jean du Moroni; Venezuela:
Salto de las Acadencias.

Enchophora uniformis, n. sp.

Fig. 72

Length. — Male 24-26 mm, female 25-26
mm.

Ground color yellow to greenish yellow.
Head process scimitar-shaped, shorter than
frons below process. Hind wing white with
apical third yellowish. Tegmen with margin
darker behind stigmal line; this margin with
eight spots that may be indicators of waxy
tufts. Distinctive markings, brown or black:
apex of supraocular flange, three or more
dashes along clavus, spots along stigmal line
(these are variable; one specimen lacks them
all).

This species has a smaller process than
viridipennis and prasina, the only two other
species with green or yellow tegmen. It lacks
the white costal margin of prasina and the
yellow-orange base of the hind wing of viri-
dipennis.

Distribution. — Holotype (male), allotype
(female), and 1 paratype (female): Peru: Iqui-
tos, F6062, H. Bassler Golln. Ace. 33591
(AMNH). Four other paratypes: PERU: Chau-
chamayo [sic], from W. F. H. Rosenberg
(NCU, male); Jauja Prov. Satipo, Sept. 1946,
A. Mailer Coll., Frank Johnson Donor
(AMNH, female).; Dept. Huanuco, Vic.
Tingo Maria, Jungle, 670 m, XII- 1939, Felix
Woytkowski (AMNH, male); Iquitos, Rio
Itaya, Herbert Osborn Colin (OSU, female).

Enchophora viridipennis Spinola

Fig. 74

Enchophora viridipennis Spinola 1839:225, pi. 11, Fig. 2.
Type repository: NHMV.

Enchophora erninenta Schmidt 1909:187. Type reposi-
tory: IZW. New synonymy.

Length. — Male 22 mm.

Ground color yellow brown (given as green
in original description). Head process scimi-
tar-shaped, shorter than most other species
(Fig. 74). Hind wings yellow-orange at base,
posteriorly white with ochre veins. More than
10 waxy tufts present along slightly infuscate
margin of tegmen.

This species may be separated from all
other Enchophora but uniformis by the col-
oration of the hind wings. In the specimen
examined the distinctive color markings are

156

Great Basin Naturalist Memoirs

No. 12

the shiny, black apex of the head process, a
black spot on each side of the frons before the
eye, five very small, red marks on the tegmen,
red longitudinal lines on the clypeus, a diago-
nal, red, marginal band on each side of the
pronotum, and the browner wing margin.
Schmidt (1909) described the clypeus as hav-
ing blood-red longitudinal stripes, the apex of
the clypeus, the pronotum and scutellum, the
legs, and the five spots on the tegmen blood
red. The specimen at hand has fainter col-
oration. Spinola (1839) said that the spots on
the tegmen are black. I did not have a speci-
men to compare with either of these types
when I visited their respective museums; so, I
am placing them in synonymy by utilizing
their descriptions and illustrations and my
notes.

Distribution. — Holotype: Brazil: Pebas
(eminenta) , locality unknown {viridipennis). I
have seen one specimen from BRAZIL: Jacarea-
canga, collected in February.

Enhydria Walker

Figs. 36-39

Enhydria Walker 1858:44. Type-species: Dichoptera

tesellata Walker, by monotypy.
Ulubra Stal 1866:133. Type-species: U. hrachialis Stal,

by monotypy. Synonymized by Stal 1870:286.

Ulubra resurrected by O Brien 1985:662. [Error]

Netc synompny.

Medium-sized, brown insects with trans-
parent tegmina, 19-22 mm long. Head pro-
cess recurved, laterally compressed, gradu-
ally narrowing to apex. All five ventral carina
and two dorsolateral carina present, com-
plete; dorsal median carina present at base of
process, extending as far as dorsal notch. Pre-
ocular flange horizontal, small. Pronotum
with white nodules. Tegmen with short spines
along veins on upper surface, M not pectinate;
cross-veins along costal margin dense, diago-
nal; other cross-veins not dense, about same
diameter as longitudinal veins; most cells
rectangular. Female with eighth and ninth
abdominal tergites subequal.

This genus may be separated by the short
spines on the veins on the upper surface of the
tegmen and the transparent, brown hind
wing. The dorsal head measurement and
head/pronotal ratio were not made in this
genus because the length varies greatly with
the position of the head process.

I resurrected Ulubra as a valid genus on the

basis of illustrations and notes taken from a
specimen so identified at the British Mu-
seum. Recently, I discovered a photograph of
the type of Ulubra brachialis taken in Stock-
holm; and it is a synonym of tessellata. The
genus I resurrected as Ulubra is renamed he-
lowAS Stalubra(q. v.).

All species o( Enhydria have a similar color
pattern: brown insects with red spots on the
pronotum, black markings on the head pro-
cess, etc. They may be identified to species by
the length and shape of the head process.

Key to the Species oi Enhydria

1. Head in lateral view with distance from fronto-
clypeal suture to dorsal notch greater than
length from dorsal notch to apex, thus process
short (Figs. 37, 38) 2

— Head in lateral view with distance from fronto-
clypeal suture to dorsal notch on process less
than length from dorsal notch to apex, thus pro-
cess long (Fig. 36) longicornuta Lallemand

2(1). Venter of process black; thickness of process in
lateral view about 1/2 width of eye (Fig. 38) . . .
cicadina Gerstaecker

— Venter of process pale with brown spots, only tip
black; thickness of process in lateral view sube-
qual to width of eye (Fig. 37)

tessellata (Walker)

Enhydria cicadina Gerstaecker

Figs. 38, .39

Enhydria cicadina Gerstaecker 1895:38. Type repository:
EMAU.

Length. — Male 19.5-21.5 mm.

This species has the shortest head process
(Fig. 38).

Male genitalia. — The two pairs of inflat-
able lobes have darkened, tuberculate tips,
which the other species do not have.

Distribution. — Syntypes: Brazil: Bahia.
I have seen six other specimens collected
in December from Brazil: Caviuna, Iguazu
Falls, and Tijuco Preto.

Enhydria longicornuta Lallemand

Fig. 36

Enhydria brachialis i. longicorntita Lallemand 1960:106.

Type repository: NRS.
Enhydria longicornuta Lallemand, O Brien 1985:661.

Length— Male 20-22 mm, female 20-22
mm.

This species has the longest head process
(Fig. 36).

Male genitalia. — ^The ventral lobe in ven-
tral view has a slight median projection and is

1988

O'BRIEN: New World Fulgoridae

157

about half as wide as the claspers.

Distribution. — Holotype: Brazil. I have
seen five other specimens collected from
Brazil: Manaus, Serro do Navio, and Trin-
idad.

Enhydria tessellata (Walker)

Fig. 37

Dichoptera tessellata Walker 185L305. Tvpe repositorv:

BMNH.
Enhydria tessellata (Walker), Walker 1858:44.
Enchophora brachialis Stal 1862: L Type repository:

NHS. New sijnonytmj.
Enhydria brachialis (Stal), da Costa Lima 1935:497.

Length. — Male 19-20 mm, female 21-22
mm.

This species has a medium-length head pro-
cess (Fig. 37).

Male genitalia. — Ventral lobe of the
aedeagus with median emargination, ventral
lobe about as wide as claspers.

Distribution. — Holotype: Brazil: Rio de
Janeiro (brachialis male); Para {tessellata fe-
male). I have seen 11 other specimens col-
lected in January, March, June, and October
from Brazil: Corcovado, Faz Morelandia,
Corupa, and Para; PERU: Tingo Maria.

Fulgora Linnaeus

Figs. 77-92

Fulgora Linnaeus 1767:703. Type-species: F. laternaria
(Linnaeus), subsequent designation by Interna-
tional Commission on Zoological Nomenclature,
Opinion 322 (1954:187).

Large insects, 65-105 mm in length, yel-
lowish brown mottled with black markings
and white, waxy areas, head .25 to .28 length
of insect. Head process porrect, terete, in-
flated, resembling a peanut from above and an
alligator head, including false eye spots, in
lateral view. Preocular flange large, horizon-
tal. Tegmen with M not pectinate, most cross-
veins forming many-sided, almost round
cells. Female with abdominal tergites eight
and nine subequal in length.

At the museum in Sao Paulo, 1 examined
the specimens used by da Fonseca (1926) in
his revision; these did not fit his key. B. V.
Ridout (personal communication) examined
previously used characters and several new
features, including male genitalia and the re-
sults of morphometric analyses, and found the
best way to identify species in this genus was
to match specimens to the accompanying il-
lustrations of head shapes. His fllustrations

are labeled lectotype except for la7npetis
(holotype), crocodilia, and lucifera. I give him
full credit for this section and am convinced
that this is the best that can be done at this
time. Any errors are mine.

Ridout suggested that species with narrow
heads might live in dry forests, the others in
rain forests. Janzen (1983) described the be-
havior of F. laternaria, including at least 100
specimens seen resting on Hymenaea cour-
haril or guapinol, but questioned whether
this plant is the host or only host, as it does not
extend throughout the range of the insect spe-
cies.

Fulgora castresii Guerin-Meneville

Figs. 81, 82

Fulgora castresii Guerin-MeneviWe 1837:3, pis. 173, 174,
Figs. 3, 4.

Distribution. — Type: Mexico. I have seen
10 specimens collected from July through
September from MEXICO: Hustusco, Nuevo
X-Can [sic], Tamazunchale; Peru: Tingo
Maria: Panama: Barro Colorado Island.

Fulgora cearensis Forseca

Figs. 87, 88

Laternaria cearensis Fonseca 1932:3, Figs. 1, 2, 3h.
Fulgora cearensis Fonseca, Metcalf 1947:219.

Distribution. — ^Type: Brazil. I have seen
two specimens from Trinidad : Curepe and
Maraval, collected in September.

Fulgora crocodilia
Brailovsky & Beutelspacher

Figs. 85, 86

Fulgora crocodilia Brailovsky & Beutelspacher 1978:176,
pi. 3.

Distribution. — Type: Mexico. I have
seen three specimens collected from MEXICO:
Chamela, in September and November.

Fulgora graciliceps Blanchard

Figs. 83, 84

Fulgora graciliceps Blanchard 1849:pl. 2, Fig. 1.
Laternaria orthocephala Fonseca 1926:493, pi. 6, Figs. 1,

2. New synonymy, Ridout.
Fulgora orthocephala Fonseca, da Costa Lima 1942:42.

Distribution. — Type: Brazil (orthoceph-
ala, graciliceps unknown). I have seen three
specimens from Bolivia: Requena and
Warnes, and PERU: Tingo Maria, collected in
November.

158

Great Basin Naturalist Memoirs

No. 12

14.2mm

I \ nn 19. Vuloin-ii Itiuwctis Burnieister;

Figs. 77-101. Head, dorsal view (odd numbers)^ ^t^J^;;;: ^8^4 T^^S Bbn'wl; SS^SG, F. crocodiUa
79-80, F.iaternam(L.); 81-82, F.casfrm.Guenn-Menevlle^ 8.3 M j^ ^^_y^ ^ riofirandensts

BraiWsky & Beutelspacher; 87-88, F. -^Xvan Xzt 94 1 voi^^ 95, A. u^ankinsi OBnen; 96^A.

Fonseca. Hind wings oi Arnycle:93, A. '^""f^ll"^^^^^^^^^ 99 a brevis O'Brien; 100, A. f.i»Ki«iror««e K & K,

pinyonae K & K; 97, A. vernalis Manee; 98, A. grandts O Bnen, 99, A.
101, A. amabilis (Westwood).

1988

OBrien: New World Fulgoridae

159

Fulgora lampetis Burmeister

Figs. 77, 78

Fulgora lampetis Burmeister 1845:3.

Distribution. — Type: Brazil. I have seen
11 specimens collected throughout the year
from Bolivia: La Cordillera Santa Cruz;
Peru: Tingo Maria; Brazil: Obidos, Sinop;
Panama: Barro Colorado Island; Costa Rica:
Sarapiqui; Nicaragua: Leon; Honduras: La
Ceiba.

Fulgora laternaria (Linnaeus)
Figs. 79, 80

Cicada laternaria Linnaeus 1758:434.
Fulgora laternaria Linnaeus, Linnaeus 1767:703.
Fulgora servillei Spinola 1839:214. New synonymy. Rid-
out.

Distribution. — Type: Tropical America
{laternaria); Brazil {servillei). I have seen
live specimens collected in January, and April
through July in Panama: Barro Colorado Is-
land; Honduras: Armenia, La Ceiba; Mex-
ico: LosTuxtlas.

Fulgora lucifera Germar

Figs. 89, 90

Fulgora lucifera Germar 1821:100.

Fulgora mitrii Burmeister 1867;xxiii, Berg 1879:178.

Distribution. — ^Type: Brazil. I have seen
16 specimens collected from November
through February from Argentina: Abra
Grande Iran; Bolivia: Coipa, El Torno, Mora
Abadolz., Montero, Santa Cruz, andWarnes.

Fulgora riograndensis Fonseca
Figs. 91, 92

Laternaria servillei riograndensis Fonseca 1926:486, pi.

IV, Fig. 1.

Laternaria riograndensis Monte 1932:22.

Fulgora riograndensis Fonseca, Metcalf 1947:232.

Distribution. — Type: Brazil. I have seen
one specimen from Panama: Barro Colorado
Island, collected in June.

Odontoptera Carreno
Figs. 44, 116, 117

Odontoptera Carreno 1841:275. Type species: O.
spectabilis Carreno, by monotypy.

Medium-sized, green or brown insects,
27-37 mm long, head 1/6 to 1/3 length of
insect. Head process porrect, terete, elongate
triangular; apex bent dorsad; lateral carinae of
dorsum and venter and pleural carinae all visi-

ble. Preocular flange small, diagonal or hori-
zontal. Ventropleural carina of pronotum
present, dorsopleural carina absent, repre-
sented by low ridge. Tegmen with apex trun-
cate; anal angle angulate; dark spot along stig-
mal line; M greatly expanded, 16-19
branches at level of stigmal line; cross-veins
incomplete in corium, forming rectangular
cells in membrane. Female with eighth and
ninth abdominal tergites subequal in length.

Cathedra serrata and the two species of this
genus are the only Fulgoridae with head pro-
cesses which I have never seen misidentified.

Key to the Species oi Odontoptera

1. Head about 1.5 times as long as pro- and
mesonota combined; tegmen green with nar-
row, apical, brown band; hind wing green with
brown and orange markings .... carrenoi Signoret

— Head almost as long as pro- and mesonota com-
bined; tegmen orange-brown basally, brown be-
hind pale, thin, nodal band; hind wing white at
base, brown apicalUy spectabilis Carreno

Odontoptera carrenoi Signoret

Figs. 44, 117

Odontoptera carrenoi Signoret 1849:178, pi. 6, Fig. 4.
Type repository: NHMV.

Length. — Male 27-28 mm, female 30-33
mm; head length 9.7 mm; ratio, head/prono-
tum5.4.

Ground color green with brown apical band
on tegmen, round, black spot surrounded by
white line at apex of clavus on stigmal line
(Fig. 44). Hind wing green with two black
spots near anal angle, spots surrounded by
orange area.

This species is green with a more elongate
head than the brown spectabilis.

Distribution. — Holotype: locality un-
known. 1 have seen 26 other specimens, col-
lected May to October and February, from:
Brazil: Sinop; French Guiana: Montague
des Singes near Kourou; 67 km S Cayenne;
Panama: Barro Colorado Island, Santa Rita
Arriba, Parque Nacional Soberania Frijolito;
Costarica: La Selva; Mexico: Los Tuxtlas.

Odontoptera spectabilis Carreno

Fig. 116

Odontoptera spectabilis Carreno 1841:277, pi. 5, Fig. 2.
Type repository: ZMHB.

Length. — Female 36-37 mm; head length
5.8 mm; ratio, head/pronotum 2.3.

160

Great Basin Naturalist Memoirs

No. 12

Ground color yellow-orange on head, tho-
rax, and apical part of tegmina, tegmen be-
hind stigmal line brown. Head, thorax, and
tegmen with lateral, longitudinal, brown
band extending half length of costa. Hind
wing white, black along apical margins, tinged
with orange at base.

This species is brown and has a shorter head
than the green carrenoi.

Distribution. — Type: America (?). I have
seen four other specimens, collected from
January to March, from Brazil: Boraceia
Field Station and Corupa.

Phrictus Spinola

Figs. 102-115

Phrictus Spinola 1839:216. Type-specie.s: Fulgora di-
adema Linnaeus by nionotypy.

Large, reddish brown or yellowish brown
insects, rarely yellow and green, 29-53 mm;
head 1/8 to 1/4 length of insect. Head process
porrect, terete, usually enlarged into trifur-
cate apex (Figs. 102-115); dorsolateral carinae
each with two pairs of spines, ventrolateral
carinae each produced into flange near fronto-
clypeal suture and into small, dull spine at
junction of frons and clypeus. Preocular flange
horizontal, small; supra- and postocular
flanges produced into strong spines. Tegmen
with M not pectinate, cross-veins reticulate in
corium forming square cells behind stigmal
line. Female with ninth tergite medially
shorter than eighth.

These beautiful insects with a name that
means "horrible" or "terrible" may be identi-
fied by the shape of the apex of the head
process and the waxy tufts on it, the color
pattern of the tegmina, and the color of the
hind wings. Galdwell (1945) also used the me-
dian notch in the caudal margin of the prono-
tum and variation in the furcation of the me-
dian carina around the notch, which I found
difficult to quantify. The female first valvulae,
as he illustrated, also serve to identify difficult
specimens. The apex and venter of the head
process are usually reddish orange or reddish
brown with a white stripe down the dorsum
reaching the apex of the pronotum. There are
several color patterns on the tegmina which
will be discussed under species descriptions. I
have illustrated the head process with the
insect at approximately a 45-degree angle in
order to provide a good view of the apex.

Because this makes the length of the head and
process difficult to determine, I have included
a head length and head/pronotum ratio.

Key to the Species of Phrictus

1. Head process flattened apically, transversely

arcuate, lacking definite apical teeth (Fig.
1 12); base of hind wing yellow with transverse

bands of brown maculae

auromaculatus Distant

— Head process with 3 to 5 apical teeth (Figs.
106-111, 113, 114); hind wing lacking trans-
verse bands of brown maculae 2

2(1). Head process with 5 apical teeth (Fig. 104)

quinquepartitus Distant

— Head process with 3 apical teeth 3

3(2). Hind wing with large (ca 2 mm), hyaline,
apical spots, visible from below with wing
unspread 4

— Hind wing without hyaline, apical spots, but
small pruinose areas sometimes present, visi-
ble when wings are spread 5

4(3). Hind wings yellow or orange at base; head
process with 3 black spots dorsally just behind
apical area (Fig. 113) ... . xanthopterus Schmidt

— Hind wing red at base; head process with no
black markings near apex, but diamond-
shaped, red area in same position (Fig. 107)
oceUatus Signoret

5(3). Basal area of hind wing yellow or yellow-
orange moebiusi Schmidt

— Basal area of hind wing red or reddish orange

6(5). Tegmen with continuous, broad, yellow,

transverse fasciae present 7

— Tegmen lacking transverse fascia, or, if
present, interrupted medially 8

7(6). Tegmen brown, transverse fascia yellow only

diligens, n. sp.

— Tegmen scarlet with green veins, anterior
edge of fascia partially bordered with black
tripartitus Metcalf

8(6). Transverse fascia absent, apical trident of
head process 2/3 as wide as distance between

supraocular spines (Fig. 108)

delicatus, n. sp.

— Transverse fascia interrupted medially; apical
trident of head process wider (Figs. 105, 106,
109, 110) 9

9(8). Head process with apical teeth broader than
head (Fig. 105); hind wing with black or fus-
cous area covering apical three-lourths

recalls Caldwell

— Head process with apical teeth not wider than
head (Figs. 106, 109, 110); hind wing with
black or fuscous areas covering apical third . 10

10(9). Tegmen green in basal two-thirds, with a few
small, roinid, red or orange maculae; apical

third with large, brown spots

hoffmannsi Schmidt

1988

OBrien: New World Fulgoridae

161

Figs. 102-117. Head and pronotum, dorsal view, drawn with head process at 45-degree angle so that the apex is
horizontal; 102, Phrictus diadema L.; 103, P. tripartitus Metcalf; 104, P. quinquepartitus Distant; 105, P. regalis
Caldwell; 106, P. punctatus Caldwell; 107, P. ocellatus Signoret; 108, P. delicatus O'Brien; 109, P. minutacanthis
Caldwell; 110, P. hoffmansi Schmidt; 111, P. moebiusi Schmidt; 112, P. auromactdatus Distant; 113, P. xanthopterus
Schmidt; 114, P. diligens O'Brien; 115a, P. diadema (L.) and 115b, P. triportitus Metcalf (head, lateral view); 116,
Cathedra spectabilis Carreno; 117, O. carrenoi Signoret.

162

Great Basin Naturalist Memoirs

No. 12

— Tegmen brown or reddish brown, sometimes
with yellow spots 11

11(10). Tegmen some shade of reddish brown, with
pink, calloused areas present, especially
basally; head process toothed on ventral sur-
face (Fig. 115a) diadeina (Linnaeus)

— Tegmen brown, maculate with yellow; head
process not toothed ventrally 12

12(1 1). Combined length of head process and head as
long as pronotum; expanse of trifurcate apex
equal to distance between ocular spines (Fig.
109) minutacanthis Caldwell

— Combined length of head process and head
longer than pronotum; expanse of apex much
greater than distance between ocular spines
(Fig. 106) punctatus Caldwell

Phrictus auromaculatus Distant

Fig. 112

Phrictus auromaculatus Distant 1905:672. Type reposi-
tory: BMNH.

Phrictus notatus Lallemand 1931:188. Type repository:
FSAG, NRS. New synonymy.

Length. — Male 29-32 mm, female 34-35
mm; head length 3.7 mm; ratio, head/prono-
tum 1.0.

Head process semicircular at apex, not di-
vided into teeth, orange-red; three round,
waxy plates on each side (Fig. 112). Tegmen
evenly pale brown mottled with yellow, veins
dark brown in corium, no transverse fascia.
Hind wings yellow at base, brown posteriorly;
two maculate, brown bands from anal area
running into yellow area.

This species is easily separated by the apex
of the head not being divided into spinelike
segments. It may be verified by the yellow
base of the hind wings and the evenly colored
tegmina.

Distribution. — Syntypes: Bolivia (auro-
maculatus), Ecuador (notatus). I have seen
five other specimens collected from Bolivia:
Santa Cruz; Prov. Sara [sic], 450 m; Brazil:
Sinop; Peru: Jauja Prov., Satipo.

Phrictus delicatus, n. sp.

Fig. 108

Length. — Female 36 mm; head length 3.7
mm; ratio, head/pronotum 1.3.

Head process tripartite, lateral portions di-
rected anterad, not laterad, smaller than that
oi minutacanthis; head pale brown (Fig. 108).
Mesonotum with three pairs of dark spots (pin
is placed through apex; there may or may not
be one there). Tegmen evenly colored, no

fascia. Hind wings red basally, posteriorly
pale brown.

This species may be identified by the pale
brown, narrow head process, without a white
dorsal stripe and red venter and apex. The
pale apex of the hind wings separates this
species from all but moebiusi. The evenly col-
ored tegmen, broader and less rooflike than
most of the species, is similar to that of auro-
maculatus and ocellatus.

HoLOTYPE (female). — Brazil: Amazonas,
S. Paulo de Olivenca, Vn-32, A. Mailer
Colin., Frank Johnson Donor (AMNH). The
printed label says S. P. Olivenca/ Amazonas/
Brasil Vn-32. The handwritten label under-
neath gives the more complete data.

Phrictus diadema (Linnaeus)
Figs. 102, 115a.

Fulgora diadema Linnaeus 1767:703. Type repository:

UZIU.
Phrictus diadema (Linnaeus), Spinola 1839: 219.

Length. — Male 38 mm, female 45 mm;
head length 8 mm; ratio, head/pronotum 2.0.

Apex of head process trifurcate, three or
four pairs of lateral, waxy plates and one me-
dian oval (Fig. 102). Ventral margin of head
process in lateral view sinuate. Hind wings
red at base. Tegmina mottled with red and
brown in large splotches, transverse, yellow-
ish fascia narrow, not bordered anteriorly in
black.

This species most closely resembles regalis
in the mottled color pattern of the tegmen and
pale, transverse fascia and the sinuate ventral
margin of the head process in lateral view, but
regalis has a much wider, trifurcate apex on
the head process, exceeding the distance be-
tween the tips of the supraocular spines (Fig.
105). In diadema the spines and apex are
about the same width.

Distribution. — Type: India ? [error]. I
have seen two other specimens collected in
November from BRAZIL: Linhares.

Phrictus diligens, n. sp.
Fig. 114

Length. — Female 40-43 mm; head length
6 mm; ratio, head/pronotum 1.3.

Apex of head process trifurcate, narrow
band of waxy plates each side (Fig. 114). Hind
wings red at base. Tegmen evenly brown with
unbroken clear transverse fascia.

This species may be identified easily by the

1988

O'BRIEN: New World Fulgoridae

163

browii tegmen with the unbroken transverse
band. I name it diligens, which means prim,
because it is more conservative in color and
shape of head process than the other species.
It is most closely related to diadema and re-
galis in the color pattern of the tegmen.

HoLOTYPE (female). — Colombia: 1,500 ft,
Anchicaya, 26-VII-1970, J. M. Campbell
(LOB). Paratype (female): Colombia: Dept.
El Valle, Anchuaya, 200-300 mi, 27-IV-
1969, Gonin-H Balazar-S. Draachsler Rec.
(MNHP).

Phrictus hoffmannsi Schmidt

Fig. 110

Phrictus hoffmannsi Schmidt 1905:338. Type repository:
IZW.

Length. — Male 37 mm, female 48 mm;
head length 7 mm; ratio, head/pronotum 1.0.

Apex of head process trifurcate, large,
squarish, waxy plate area on each side
(Fig. 110). Hind wings red at base. Tegmina
green mottled with yellowish orange in
large splotches, clavus predominately orange,
broad, transverse, yellow fascia interrupted
by brown spots medially and toward commis-
sure, membrane yellowish brown with brown
oval spots.

This species may be distinguished easily by
its bright green and yellowish orange tegmen.

Distribution. — Holotype (female): Peru:
Chanchamayo. I have seen four other speci-
mens collected in January from Peru: Tingo
Maria; Ecuador: Guayaquil, Loja.

Phrictus minutacanthis Caldwell
Fig. 109

Phrictus jninutacanthis CaldweW 1945:180, pi. 7, Figs. 2,
16, pi. 8, Fig. 2, pi. 9. Type repository: USNM.

Length. — Female 36 mm; head length 4.8
mm; ratio, head/pronotum 1.5.

Apex of head process trifurcate, small; waxy
plate on each side; pale red, midline near apex
with reddish triangle (Fig. 109). Hind wings
red at base. Tegmina light brown evenly mot-
tled with yellow, transverse yellowish fascia
indistinct, broken.

This species may be identified by its small
head. It resembles xanthopterus and punc-
tatus most closely in the light brown tegmen
with yellow markings, and ocellatus and punc-
tatus in having the red subapical triangle on
the pale dorsum of the head process.

Distribution. — Holotype: Peru: Chaqui-
mayo. I have seen only the type.

Phrictus moebiusi Schmidt
Fig. Ill

Phrictus moebiusi Sclimiclt 1905:335. Type repository:

ZMHB.
Phrictus sordidus Caldwell 1945:180, pi. 7, Figs. 3, 20,

pi. 8, Figs. 1, 13, pi. 9. Type repository: USNM.

New synonymy.

Length — Male 40 mm, female 43-48 mm;
head length 6 mm; head/pronotal ratio 1.2.

Apex of head process trifurcate, large, waxy
plate on apex (Fig. 111). Hind wings yellow at
base, posterior pale brown. Tegmina brown
evenly mottled with yellow, transverse fascia
pale, broad, interrupted by two brown spots
in middle and near commissural margin.

This is one of three species with yellow
coloration at the base of the hind wing. P.
auromaculatus has a head process without
apical spines, and xanthopterus has white
spots in the posterior brown area of the hind
wing. The type of sordidus is a pale specimen
that agrees in all external characters with moe-
biusi.

Distribution. — Syntype (female): Colom-
bia: Antioquia (moebiusi); Ecuador (male
holotype sordidus). I have seen four other
specimens collected in January from "New
Granada" and Ecuador: Route Mono-Los
Banos, km 37, 2,385 m.

Phrictus ocellatus Signoret
Fig. 107

Phrictus ocellatus Signoret 1855:v. Type repository:

nhmv.

Length. — Male 35 mm, female 42 mm;
head length 6.0; ratio, head/pronotum 1.9.

Apex of head process trifurcate; large, waxy
plates on each side; red diamond and median
line along pale dorsum of process (Fig. 107).
Hind wings red at base, white spots in brown
posterior area. Tegmina evenly mottled with
pale yellow and red brown, no fascia.

Two species have hind wings with white
spots in the brown posterior area, xan-
thopterus with a yellow base and ocellatus
with a red base.

Distribution — Holotype (female): Co-
lombia: Fusagasuga. I have seen two other
specimens collected in June from VENEZUELA:
Rancho Grande.

164

Great Basin Naturalist Memoirs

No. 12

Phrictus punctatus Caldwell

Fig. 106

Phrictus punctatus Ca\dwe\\ 1940:181, pi. 7, Figs. 12, 23,
pi. 8, Fig. 6, pi. 9. Type repository: USNM.

Length. — Female 40 mm, head length 6.5
mm; ratio, head/pronotum 1.4.

Apex of head process trifurcate, with trans-
verse, waxy plates; triangular red area on mid-
line behind apex (Fig. 106). Hind wings red at
base, pale brown posteriorly. Tegmina brown
mottled with yellow in large splotches, clavus
predominantly yellow; transverse, yellowish
fascia narrow, interrupted medially; diagonal
yellow line from clavus to costal margin.

In coloration, the tegmen o( punctatus most
closely resembles xanthoptertis, while the
color of the head process resembles oceUatus
and minutacanthis.

Distribution. — Holotype (female): Pana-
ma: Bugaba. Paratype: El Volcan. I have seen
two other specimens, one without data, the
other collected from CosTA RiCA: Palo Verde.

Phrictus quinquepartitus Distant

Fig. 104

Phrictus quinquepartitus Distant 1883:24, pi. 4, Fig. 8.
Type repository: BMNH.

Length. — Male 40-42 mm, female 45-48
mm; head length 9.3 mm; ratio, head/prono-
tum 2.06.

Apex of head process trifurcate, one median
and three lateral pairs of waxy tufts in apex
(Fig. 104). Hind wings red at base. Tegmina
red with green veins; transverse, yellowish
fascia bordered anteriorly in black.

This species is closely related to tripartitus
in the color of the tegmen but differs in the
apex of the head process.

Distribution. — Syntypes: Panama: Buga-
ba, 3,000-4,000 ft; Colombia: Bogota. I have
seen nine other specimens collected in Feb-
ruary, June, August, October and December
from Panama: Barro Colorado Island, Code,
Chiriqui; CoSTA RiCA: La Selva.

Phrictus regalis Caldwell
Fig. 105

Phrictus regalis Ca\dwe\\ 1945:183, pi. 7, Figs. 7, 19, pi.
8, Fig. 8, pi. 10. Type repository; USNM.

Length. — Male 42 mm, female 49-53 mm;
head length 10; ratio, head/pronotum 2.3.

Apex of head process trifurcate, three waxy
tufts on each side, two larger ones medially.

semicircular carinate area astride median
carina (Fig. 105). Hind wings red at base.
Tegmina mottled with red and brown in large
splotches; transverse, yellowish fascia nar-
row, not bordered anteriorly in black.

This species is very similar to diadema in
the color of the tegmen, but the apex of the
head process is much wider in regalis (Figs.
105, 102).

Distribution. — Holotype (female): French
GuiANA: Maroni River, near Duserre. I have
seen three other specimens collected in
March from French Guiana: St. Jean du Ma-
roni; Suriname: Nassaugebergte.

Phrictus tripartitus Metcalf

Figs. 103, 115b

Phrictus tripartitus Metcalf 1938:365, pis. 20, 21. Type
repository: AMNH.

Length. — Male 35-40 mm, female 43-45
mm; head length 9.4 mm; ratio, head/prono-
tum 1.8.

Apex of head process trifurcate, three
round, waxy plates and one oval on each side
(Fig. 103). Head process sinuate ventrally in
lateral view (Fig. 115b). Hind wings red at
base. Tegmina red with green veins; trans-
verse, yellowish fascia bordered anteriorly in
black.

This species most closely resembles quin-
quepartitus in the color of the tegmen, but the
species may be separated by the shape of the
apex of the head process. Some specimens of
tripartitus do have five spines, but the inter-
mediate spines are smaller than those of quin-
quepartitus (Fig. 104) and the shape of the
other teeth is like that illustrated for triparti-
tus (Fig. 103).

Distribution. — Type: Belize. I have seen
21 other specimens collected from August
through October from Belize: 24 mi SE Bel-
mopan, 10-11 mi S Georgeville; Honduras:
La Ceiba; MEXICO: Nuevo X-Can [sic], San
Quintin, Yucatan.

Phrictus xanthupterus Schmidt

Fig. 113

Phrictus xanthopterus Schmidt 1910:144. Type reposi-
tory: IZN.

Length. — Female 39-41 nnn; head length
5 mm; ratio, head/pronotum 1.0.

Head process with apex trifurcate, without
waxy plates, orange-red, with three triangu-
lar, black spots behind apex (Fig. 113). Hind

1988

O'BRIEN; New World Fulgoridae

165

wings yellow at base with white spots in
brown posterior area. Tegmen brown with
yellow markings, half of clavus predominantly
yellow; transverse, yellow fascia interrupted
medially.

This species is one of the most distinctive
with the hind wings yellow at base and with
white spots in the apical brown area and the
three black, subapical spots on the head pro-
cess. The tegmen most closely resembles
punctatus.

Distribution. — Holotype (female): Ecua-
dor: Canelos. I have seen six other specimens
collected in June and July and December
from Peru: Cusco; Sta. Isabel; Route Olmoc-
Tarapoto, km 385, 1,800 m; Tingo Maria;
Ecuador: Lumbagui.

Rhabdocephala Van Duzee

Fig. 23

Rhabdocephala Van Duzee 1929; 190. Type-species; R.
brunnea Van Duzee, original designation.

Small, brown, narrow insects, 12-16 mm
long; head about 1/3 length of insect. Head
process porrect, terete, sides parallel; apex
slightly widened; dorsal lateral carinae com-
plete, ventral incomplete, present only in
middle of process. Preocular and supraocular
flanges absent. Tegmen with M not pec-
tinate, cross-veins reticulate, apical margin
angled at apex of clavus, forming angulate, not
smoothly rounded, apical margin. Ninth ab-
dominal tergite 1.5 times length of eighth.

These small insects may be separated from
other small Fulgoridae by the simple cephalic
process with no dorsolateral undulate carinae
as in Scolopsella and the head process not
dorsoventrally flattened as in Amycle.

Rhabdocephala brunnea Van Duzee

Fig. 23

Rhapdocephala brunnea Van Duzee 1929; 191. Type
repository; CAS.

Length. — Male 12-14 mm, female 14.5-
16 mm; "head 5.2 mm; ratio, head/pronotum
2.1.

Tegmen brown mottled with white. Hind
wing red in basal third, brown posteriorly.

Distribution. — Holotype (female): USA:
Arizona. I have seen 15 other specimens col-
lected from May to October from USA: ARI-
ZONA: Pima, Cochise, and Santa Rita counties;
Mexico: Sonora, 15 mi S Hermosfllo. Two

specimens were collected on Baccharis
sarothroides Gray.

Scolopsella Ball

Figs. 8-13, 24-25

Scohypsella Ball 190.5; 118. Type-species; S. reticulata
Hall by original designation.

Small, brown insects, 14-21 mm long, head
.23-. 28 length of insect. Head process por-
rect, terete, very slightly enlarged at apex;
dorsolateral carinae undulate; median and lat-
eral ventral and marginal carinae complete on
process and frons. Preocular flange indistinct.
Tegmen with M pectinate (3 branches), com-
missural margin emarginate between claval
apex and wing tip.

This genus of small, brown insects may be
separated from other similar genera by the
straight head process with dorsolateral mar-
gins crenulate or undulate and without the
apex greatly expanded. The head process is
not as unadorned as Rhabdocephala, lacks the
transverse wrinkles o( Aphrodisias , and is not
flattened as in Amycle. Diareusa specimens
are much larger insects with much shorter
heads.

Key to the Species oi Scolopsella

1. Length of anal flap of male greater than width at
apical expansion (Fig. 23); apical margin of

pygofer emarginate ventrally (Fig. 25)

reticulata Ball

— Length of anal flap of male less than width at
apical expansion (Fig. 20); apical margin of

pygofer smoothly convex (Fig. 22)

mexicana O'Brien

Scolopsella niexicana O'Brien

Figs. 8-10, 24

Scolopsella niexicana O'Brien 1985:73. Type repository;
LOB.

Length. — Male 14-15 mm, female 17-21
mm; head length 5.6 mm; ratio, head/prono-
tum 3.7.

Tegmen various shades of brown, cross-
veins and reticulations pale, some apical cells
white. Structural characteristics common to
both species given in generic description.

Scolopsella reticulata and S. mexicana may
be separated by the shape of the anal flap,
which can be seen without dissection, the
applicable characters being given in the key.
Once males are identified in this way, females
may often be associated with them because
the golden brown reticulata is slightly larger

166

Great Basin Naturalist Memoirs

No. 12

and the head process is wider, and mexicana is
darker.

Distribution. — Type: Mexico. I have
seen seven other specimens collected from
June through September from Baja Califor-
nia.

Scolopsella reticulata Ball
Figs. 11-13, 25

Scolopsella reticulata Ball 1905:118. Type repository:
CAS.

Length. — Male 14-16 mm, female 17-19
mm; head length 5 mm; ratio, head/pronotum
4.1.

The structural characters and coloration do
not differ sufficiently from S. mexicana to de-
termine the species without either using the
shape of the male anal flap or comparing the
two species as discussed above.

Distribution. — Holotype: Arizona. I have
seen other specimens collected from April to
November in Arizona: Maricopa, Pima,
Pinal, and Santa Cruz counties; California:
Joshua Tree National Momument, Riverside
or San Bernardino County.

Sinuala, n. gen.
Figs. 43, 46-48

Type-species: Sintiala stall O Brien.

Reddish brown, medium-sized insects,
19-21 mm long, head .12 to .15 length of
insect. Head process porrect, terete, en-
larged at apex; vertex truncated triangle, basal
angle rounded, supraocular lobes variable.
Frons with lateral carinae diverging anteriorly
to form process; lateral margins expanded
near frontoclypeal suture, ridge from lateral
carina to margin at this expanded area. Frons
and clypeus about in same plane, frons not
raised above clypeus; clypeus medially and
marginally carinate. Preocular flange length-
ened into pleural carina. Pronotum without
raised median carina, lateral fossettes and two
pleural carinae present; apex of mesonotum
with striate triangle, delimited by carinae, not
high ridge. Tegmen with M not pectinate,
clavus open, longitudinal veins angulate,
cross-veins variable. Fore and mid femora ex-
panded. Posterior tibiae with eight lateral
spines. Female abdominal tergite nine twice
length of eight.

This genus is similar to Aphrodisias , from
which it differs by the costal margin of the

tegmen sinuate and the head process not
transversely ridged.

Key to the Species oi Sinuala

1. Supraocular flange expanded, height equal to
width of eye; tegmen with 10 strong tubercles
tuberculata, n. sp.

— Supraocular flange smaller than above; tegmen
with tubercles evanescent 2

2(1). Vertex as long as broad (Fig. 46); hind wing red
at base stali, n. sp.

— Vertex 1.7 times longer than broad (Fig. 48);
hind wing pink at base sclimidti, n. sp.

Sinuala stali, n. sp.
Fig. 46

Length. — Male 19-20 mm; head 2.3 mm;
ratio, head/pronotum 1.4.

Head process short, vertex as long as broad.
Hind wing red at base, narrow anterior mar-
gin and broad posterior area brown, apical
angle brown with some white spots and
crossveins.

This species may be easily separated from
tuberculata, which it resembles in wing color
and head length, by the small, supraocular
flange oi stali. I name this species in honor of
Carl Stal, who provided a fine basis for generic
work in this family.

Holotype (male). — Honduras: Lago
Yojoa., 6-VI-1976, J. V. Mankins, coll. (LOB).
Paratype (male): El Salvador: Alfredo Mar-
tinez Cuestas, no further data (NCSR).

Sinuala schmidti, n. sp.

Fig. 47

Length. — Female 21.5 mm; head 3.3 mm;
ratio, head/pronotum 2.2.

Head process elongate, vertex 1.7 times as
long as broad. Hind wing pink at base, then
white; anal margin pale brown, posterior mar-
gin with cells infuscate, veins white; apical
third brown with white lines following each
cross- vein.

This species may be separated from the
others by the long head and the pale base of
the hind wings. I name this species in honor of
Edmund Schmidt, who had an excellent eye
for species and a gift for describing the most
significant characters precisely.

HoLOT\'PE (female): BELIZE: Corazol, 1 mi
NW Corozal, 12-VIII-1977. C. W. and L. B.
O'Brien and Marshall (LOB).

Sinuala tuberculata, n. sp.

Figs. 43, 48

1988

O'BRIEN: New Would Fulgoridae

167

Length. — Female 21 mm; head 2.6 mm;
ratio, head/pronotum 1.6.

Head process short, vertex with postocular
flange raised above vertex equal to width of
eye, vertex as long as broad. Hind wing red in
basal half, anal area brown, apical 2/5 brown
with some white spots and cross-veins. Five
strong tuberculations on veins in anterior half
of tegmen, five smaller in apical half.

This species may be separated easily from
the others by the strong tubercles on the teg-
men and the strong supraocular flange.

Holotype (female): Costa Rica: P. R. Uh-
ler Colin. (NCSR).

Stalubra, n. gen.
Figs. 1-3, .5, 6

Type-species: Stalubra brunnea O'Brien.

Medium-sized, brown or reddish, transpar-
ent-winged insects, 19-28 mm long, head . 14
to .19 length of insect. Head process terete,
gradually narrowing to apex, upturned at
apex. Vertex twice as long as broad, elongate
medially (Fig. 1). Lateral carinae of frons par-
allel or diverging toward head process, me-
dian carina on process and basal 2/3 of frons;
frons only very slightly raised above fronto-
clypeal suture; lateral margins of frons ex-
panded near apex; weak ridge from lateral
carina to lateral margins. Clypeus laterally
carinate. Preocular flange reduced to slight
tubercle on vertical carina joining marginal
carinae of frons and vertex (Fig. 3). Pronotum
without median carina raised, fossettes and
two pleural carinae present; mesonotal apex
not depressed or striate. Tegmen with M
pectinate (5-7 branches); clavus open; cross-
veins undulate, parallel; no small spines on
veins, only on wing margins. Posterior tibiae
with six lateral spines. Female abdominal ter-
gite nine shorter than eight.

This genus is similar to Enhydria in the
compressed, recurved head process and
translucent tegmen; however, it lacks the
hairs on the veins of the tegmen and the
dense, diagonal cross-veins along the costal
margin found in Enhydria, and the head pro-
cess is less strongly curved. It differs from
Chilobia in the vertex being elongate medi-
ally, whereas the vertex in Chilobia is broader
than long and emarginate medially (Figs. 51,
54, 60) or hidden by the recurved head pro-
cess (Fig. 57).

Lallemand's species Enhydria rufula (1966)
belongs in this genus, but I cannot tell from
his description which it is of two of the three
species I have before me. The requested type
has not yet arrived, and so these two species
will be discussed in Part II.

Stalubra brunnea, n. sp.

Figs. l-,3, 5, 6

Length. — Male 19-22 mm; head 2.6 mm;
ratio, head/pronotum 1.9.

Head short, vertex 1.5 times as long as
wide. Hind wing clear with brown veins. Teg-
men translucent with brown markings and
minute red dots along veins, apex pale brown
with white circles plus some darker brown
spots.

Holotype (male). — Brasil [sic]: Mato
Grosso, Sinop (12°31'S, 55°37'W), X-1975,
M. Alvarenga (LOB). Paratypes (25 male):
same data (LOB, MZSP); Guyana: 1 male.
Upper Mazaruni River, IX-X-1938, A. S.
Pinkus(AMNH).

ACKNOWLE DGM ENTS

It is with great pleasure that I dedicate this
paper to our Latin American colleagues and to
Paul Oman, among whose many accomplish-
ments is a paper on a U.S. fulgorid (Oman
1936).

I thank the American Philosophical Society
for a grant to study at European museums
(Grant No. 5771, Penrose Fund), without
which this work would have been impossible.
I am deeply indebted to B. V. Ridout for his
work on Fulgora. I greatly appreciate the crit-
ical reading of the manuscript by F. W. Mead,
G. J. Wibmer, and S. W. Wilson and the
sending and examining of types and paratypes
of R. C. Froeschner and N. D. Penny. I am
also grateful to the following museums for
allowing me to examine types and borrow
specimens in their collections: AMNH —
American Museum of Natural History, New
York; BMNH — British Museum (Natural His-
tory), London, England; CAS — California
Academy of Sciences, San Francisco; DEI —
Institute fiir Pflanzenschutzforschung (for-
merly Deutsches Entomologisches Institut),
Kleinmachnow, Eberswalde, East Germany;
EMAU — Zoologisches Institut und Museum,
Ernst Morits Arndt Universitat, Greifswald,
East Germanv; FMNH — Field Museum of

168

Great Basin Naturalist Memoirs

No. 12

Natural History, Chicago, Illinois; FSAG —
Faculte des Sciences Agronomiques, Gem-
bloux, Belgium; IZW — Instytut Zoologii
PAN, Warsaw, Poland; LOB— Lois O'Brien
collection, Tallahassee, Florida; MCZ —
Museum of Comparative Zoology, Harvard
University, Cambridge, Massachusetts;
MHNP — Museum National d'Histoire Na-
turelle, Paris, France; MRSN — Museo Re-
gionale di Scienze Naturali, Turin, Italy;
MZF — Museo Zoologico dell' Universita de-
gli Studie di Firenze, Florence, Italy;
MZSP — Muzeo de Zoologia, Universidade de
Sao Paulo, Brazil; NCSU— North Carolina
State University, Raleigh; NHMV — Natur-
historisches Museum, Vienna, Austria;
NRS — Naturhistoriska Riksmuseet, Stock-
holm, Sweden; OSU — Ohio State University,
Columbus; UCB — University of California,
Berkeley; UMO — University Museum, Ox-
ford University, Oxford, England; UNAM —
Instituto de Biologia and Museo de Zoologia,
Universidad Nacional Autonoma de Mexico,
Mexico City, D.F.; USNM— United States
National Museum, Washington, D.C.;
UZIL — Universitets Zoologiska Institut,
Lund, Sweden; UZIU — Universitets Zoolo-
giska Institut, Uppsala, Sweden; ZMHB —
Zoologisches Museum, Humboldt Universi-
tat. East Berlin, East Germany; ZMUC-
Zoologisk Museum, Universitets Copen-
hagen, Denmark; ZSBS — Zoologische Samm-
lungen des Bayerischen Staates, Munich,
East Germany.

Literature on New World Fulgoridae
Since Metcalfs Catalog

(a complete bibliography will be included with the
final part.)

Brailovsky, a., H. and C. R. Beutelspacher B 1978.
Una nueva especie de Fulgora Linneo (Ho-
moptera: Fulgoridae) de Mexico. An. Inst. Biol.
Univ. Nal. Auton. Mexico, Ser. Zool. 49:
175-182.

Dlabola, J , AND F Zayas 1980. Eine neue Fulgoride der
Gattung javiaicastes von Cuba (Homoptera,
Auchenorrhyncha). Acta Faun. Entomol. Mus.
Nat. Pragae 16: 83-84.

Emelyanov, A. F 1979. The problem of the family dis-
tinction between the Fulgoridae and the Dic-
tyopharidae (Homoptera, Auchenorrhyncha). In:
Phylogeny and systematics of insects. Acad. Sci.
USSR, Proc. Zool. Inst. 82: 3-22.

Fennah, R C , and J. C M. Carvalho. 1963. On a new
genus and species of fulgorid from Brazil (Ho-

moptera, Lystrini). An. Acad. Brasileira Cienc.
35: 459-461.

Hemming, F 1955. Validation, under the plenary pow-
ers, of the generic name: "Fulgora" Linnaeus,
1767 (Class Insecta, Order Hemiptera) and desig-
nation for the genus so named of a type species in
harmony with current nomenclatorial practice.
Opinion 322. International Commission on Zo-
ological Nomenclature. Opin. & Declar. ICZN.
9(13): 185-208.

Horn. W., and I Kahle 1935. Uber entomologische
Sammlungen (Ein Beitrag zur Geschichte der En-
tomo-Museologie). Ent. Beih. Berlin-Dahlem 2:
1-540.

Janzen. D H , andC L Hogue 1983. Fulgora late rnaria
(machacha, peanut-head bug, lantern fly.) In:
Costa Rican natural history. University of Chicago
Press, Chicago, Illinois, xi + 816 pp.

Johnson, L. K , and R B Foster 1986. Associations of
large Homoptera (Fulgoridae and Cicadidae) and
trees in a tropical forest. J. Kansas Entomol. Soc.
59: 41.5-422.

Knull, D J . and J. N Knull 1947. Two new Amycle
(Homoptera: Fulgoridae). Ann. Entomol. Soc.
Amer. 60: 397-400.

Kramer, J P 1978. Taxonomic study of the American
planthopper genus Cyrpoptus (Homoptera: Ful-
goroidea: Fulgoridae). Proc. Biol. Soc. Washing-
ton 91: 303-335.

Lallemand, V 1956. Contribution a I'etude des Fulgori-
dae (Hemiptera) (Ire. note). Bull. Inst. R. Sci.
Nat. Belgique. 32: 1-7.

1959. Description de nouvelles especes de Ful-

gorides d'Asie et d'Afrique. Zool. Meded. 36:
267-272 [no African species, five New World].

1960. De quibusdam Fulgoris. Ent. Mitt. 24:

101-107.

1963. Nouvelles especes d'Homopteres. Bull.

Inst. R. Sci. Nat. Belgique. 39: 1-5.

1966. Fulgorides nouveaux du Musee Zoologique

de Berlin et de ma collection. Bull. Recher.
Agron. Gembloux. N. S. 1: 51-54.

Metcalf, Z p. 1947 General catalogue of the Hemiptera.
Ease. IV. Fulgoroidea, Part 9. Fulgoridae. Smith
College, Northampton, Massachusetts. 280 pp.

Nast, J 1950. New genera and species of Neotropical
Fulgoridae in the collection of the British Museum
(Homoptera). Ann. Mus. Zool. Polonici 14: 167-
175.

1951. Some remarks on Neotropical Fulgoridae

with descriptions of new genera and species (Ho-
moptera). Ann. Mus. Zool. Polonici 14: 267-279,
3 plates.

O Brien, L. B 1985. New synonymies and combinations
in New World Fulgoroidea (Achilidae, Del-
phacidae, Flatidae, Fulgoridae: Homoptera).
Ann. Entomol. Soc. Amer. 78: 657-662.

1986. Five new species of Fulgoroidea (Ho-
moptera) from the western United States and
Mexico. Southwestern Entomol. 11:67-74.

O Brien, L B , and S W Wilson 1985. Planthopper
systematics and external morphology. Pages
61-102 in L. R. Nault and J. G. Rodriguez, eds.,
The leafhoppers and planthoppers. Wiley and
Sons, Inc.

1988

O'BRIEN: New World Ful(;oridae

169

RiDOUT. B V 1983. Structure, form and function of the
lantern fly head process {Fulgora laternaria Linn.)
Unpublished dissertation, Birheck C'ollege.

Wilson, S W , and L B O Brikn 1986. Descriptions of
nymphs of Itzalana suhinaculata Schmidt (Ho-
moptera: Fulgoridae), a species new to the United
States. Great Lakes Entomol. 19: 101-105.

1987. A survey of planthopper pests of economi-
cally important plants (Homoptera: Fulgoroidea).
Pages 343-360 in Proc. Second Annual Workshop
on Leafhoppers and Planthoppers of Economic
Importance, held Provo, Utah, USA, 28 July-1
August 1986. Commonwealth Inst. Entomol.,
London.

ZiMSEN, E 1964. The type material of 1. C. Fabricius.
Munksgaard, Copenhagen. 280 pp.

List OF Species

(Countries are listed from south to north, west before
east.)

imitatrix Ossiannilsson

Venezuela, Panama,
Costa Rica, Belize, Mexico
Peru

Amerzanna O Brien, new genus

peruana O'Brien

Peru

Amycle Stal

aniahilis (Westwood)

Mexico

brevis O'Brien

Mexico

grandis O Brien

Mexico

mankinsi O Brien

Honduras

pinyonae Knull & Knull

USA:CA, AZ, NM

saxatilis Van Duzee

USA:CA

sodalis Stal

Mexico, El Salvador

tumacacoriae Knull & Knull

USA: TX, AZ

vernalis Manaee

USA: NC, LA

Aphrodisias Kirkaldy

cacica (Stal)

Mexico

shaman O'Brien

Mexico

Artacie Stal

dufourii (Signoret),

resurrected name

Fr. Guiana, Guyana

haemoptera (Perty)

Brazil

Cathedra Kirkaldy

serrata (Fabricius)

Bolivia, Peru, Brazil,

Fr. Guiana, Suriname,
Colombia, Panama, Costa Rica
Chilobia Stal
= Ecuadoria Distant, new synonymy

cinxia Stal Venezuela

dichopteroides (Distant), (Ecuadoria),

new combination Ecuador

silena Stal Ecuador

sinaragdina (Walker) Venezuela

Copidocephala Stal

guttata (White) Panama, Costa Rica,

Guatemala, Mexico
merula Distant Colombia

= melanoptera (Schmidt), new synonymy
viridiguttata Stal Columbia, Panama, Costa Rica,

Honduras, Mexico
= ornanda (Distant), new synonymy
Diareusa Walker

annularis (Olivier) Peru, Brazil, Fr. Guiana,

Suriname, Guyana
conspersa Schmidt Ecuador, Colombia

= dahli Ossiannilsson, new synonymy

keinneri Ossiannilsson

Ecuadoria, sec Chilobia

Encho})hora Spinola

atonuiria Brancsik, see Cornelia,

teste Lallcmand 1959:88
dufourii Signoret, see Artacie,

teste O'Brien 1985:661
ensifera (Germar), to inccrtae sedis
maculata O'Brien Peru

nigroinaculata Distant Bolivia, Peru, Ecuador,

Brazil
= nigrolimbata Lallcmand

pallidipunctata Lallcmand Bolivia, Peru

prasina Gerstaecker Bolivia, Peru, Panama,

Costa Rica
pyrrhocrypta Walker Brazil, Guyana, Venezuela

recurva (Olivier) Suriname, Brazil

= bohe7nani Stal, new synonymy

rosacea Distant Panama, Costa Rica, Nicaragua

sanguinea Distant Ecuador, Colombia, Panama,

Costa Rica, Nicaragua, Guatemala
=florens Distant, new synonymy
= longirostris Distant, new synonymy
stillifera (Stal) Panama, Costa Rica, Honduras,

Guyana, Belize, Mexico
subviridis Distant Panama, Costa Rica

= subviridisv ar. distanti Metcalf

tuba (Germar) Brazil

tuberctdata (Olivier) Brazil, Fr. Guiana, Suriname,

Venezuela
= tuberculata vm. fuscoinaculata Lallcmand
= parvipennis Walker, new synonymy
unifonnis O Brien Peru

viridipennis Spinola Brazil

= eminenta Schmidt, new synonymy

Enhydria Walker

= Ulubra Stal, new synonymy

cicadina Gerstaecker Brazil

ferruginea (Walker) to Hydriena (Dictyophariadae)
longicornuta Lallcmand Brazil, Trinidad

tessellata (Walker) Peru, Brazil

= brachialis (Stal), new synonymy

Fulgora L. (New synonymies in this genus are attributed

to B. V. Ridout)

caerulescens Olivier, to incertae sedis
castresii Guerin-Meneville Peru, Panama, Mexico
cearensis (Fonseca) Brazil, Trinidad

crocodilia Brailovsky & Beutelspacher Mexico

graciliceps Blanchard Peru, Bolivia, Brazil

= orthocephala (Fonseca), new synonymy, Ridout
lampetis Burmeister Bolivia, Peru, Brazil,

Panama, Costa Rica, Nicaragua, Honduras
laternaria (Linne) Brazil, Panama, Honduras,

Mexico
= servillei Spinola, new synonymy, Ridout
lucifera Germar Argentina, Bolivia, Brazil

riograndensis (Fonseca) Brazil

Odontoptera Carreno

carrenoi Signoret Brazil, Fr. Guiana, Guyana,

Panama, Costa Rica, Guatemala, Mexico

spectabilis Carreno Brazil

Phrictus Spinola

auromaculatus Distant Bolivia, Peru, Ecuador,

Brazil

170

Great Basin Naturalist Memoirs

No. 12

= notatus Lallemand, new synonymy
delicatus O'Brien Brazil

diadema (Linnaeus) Brazil, Fr. Guiana, Suriname

= diadema var. walkeri Metcalf

diligens O'Brien Colombia

hoffmannsi Schmidt Peru, Ecuador

minutacanthis Caldwell Peru

moebiusi Schmidt Colombia

= sordidus Caldwell, new synonymy
ocellatus Signoret Venezuela, Colombia

punctatus Caldwell Panama

quinquepartitus Distant Colombia, Panama,

Costa Rica
regalis Caldwell Fr. Guiana, Suriname

tripartitus Metcalf Panama, Honduras, Belize

xanthopterus Schmidt Peru, Ecuador

Rhabdocephala Van Duzee

brunnea Van Duzee USA; AZ, Mexico

Scolopsella Ball

mexicana O'Brien Mexico

reticulata Ball USA: AZ, CA

Sinuala O'Brien, new genus

schmidti O'Brien Belize

stali O'Brien Honduras, El Salvador

tuberculata O'Brien Costa Rica

Stalubra O'Brien, new genus

brunnea O'Brien Brazil, Guyana
ruftda (Lallemand), (Enhydria),

new combination Brazil

A NUMERICAL TAXONOMIG ANALYSIS OF INTERSPECIFIC MORPHOLOGICAL

DIFFERENCES IN TWO CLOSELY RELATED SPECIES OF CICADA

(HOMOPTERA, CICADIDAE) IN PORTUGAL

J. A. Quartan'

Abstract — Cicada orni Linnaeus is among the most common and widespread cicadas in Portugal, and, unless a
critical study of the male genitalia is made, it is easily confused with the much less widely distributed C. barhara
lusitanica Boulard. These species are morphologically very similar and sometimes diflkult to separate using existing
keys. This study attempts to test the discriminating capabilities of numerical technicjucs commonly used for classifica-
tory purposes, as well as to discover the most effective characters to distinguish between the two species. F"or these
purposes, cluster analysis and principal component analysis were applied to a sample of 64 male specimens character-
ized by 40 characters (33 derived from the external morphology and 7 from genitalia). In WPGMA cluster analysis,
product-moment correlations gave a better separation between these species than did taxonomic distance coefficients;
moreover, the analysis derived from the genital characters alone gave better separation than the analyses based on the
33 external characters. Principal component analysis yielded a clear, interspecific separation along the first axis. The
best characters to discriminate between males of the two species were the lengths of the pygofer (and its dorsal spine),
the tenth abdominal segment, and the appendages of the latter (which are smaller in barbara lusitanica), as well as the
width of the shaft of the aedeagus (thinner in orni).' Finally, the uniformity of the general clustering pattern resulting
from the two multivariate techniques suggests the presence of two distinct species, as also clearly indicated by
behavioral data.

Cicada orni Linnaeus is among the most
common and widespread cicadas in Portugal,
and, unless a critical study of the male geni-
talia is made, it is easily confused with the
much less widely distributed C barbara lusi-
tanica Boulard (Quartan and Fonseca 1988).
As live specimens, however, they are easily
distinguished by the male calling songs,
which are quite distinct. Oscillograms are
found in Claridge et al. (1979) and Boulard
(1982), respectively, for C. orni and C. bar-
bara lusitanica.

The two species are externally very similar
and sometimes even difficult to separate by
existing keys (e.g., Gomez-Menor 1957). In
fact, the main distinguishing character used
for their separation has been the presence in
barbara of only two spots on the cross-veins of
the forewings instead of four; however, some
specimens o( barbara lusitanica have the full
four spots as they occur in orni (Fig. 4).

Boulard (1982), when describing the Por-
tuguese form of C. barbara, which he origi-
nally named lusitanica, provided a good diag-
nosis of the genital characters of this species.
However, no detailed comparison of the two

species has been made, nor has any type of
multiple-character analysis involving the
simultaneous use of several measurements or
counts been attempted. It was felt of interest,
therefore, to see how far some common tech-
niques of numerical taxonomy would discrim-
inate between this pair of closely related spe-
cies.

This study was undertaken with two main
objectives in mind. The first was to apply
current techniques of numerical taxonomy
commonly used for class ificatory purposes
with the aim of testing their general discrimi-
nating power with respect to these two spe-
cies. The techniques chosen were a form of
hierarchical cluster analysis and principal
component analysis. It is known that apart
from the explicit use of the former, principal
component analysis can also serve as a cluster
technique of great generality and can be used
to distinguish pairs of putative morphs as in
the classical study of Temple (1968). The sec-
ond objective was to discover new characters
that might help to separate C. orni from C.
barbara.

'Depto. de Zoologiae Antropologia, Faculdade de Ciencias de Lisboa, 1200 Lisboa, Portugal.

171

172

Great Basin Naturalist Memoirs

No. 12

Material

The data on which this study is based were
taken from dried male specimens (OTUs) of 32
Cicada orni and 32 C. harhara lusitanica
(Table 1). These samples were mostly taken
by the author in Portugal: all 32 males of orni
were collected in central Portugal; an equal
number of males of harhara hisitanica were
taken in several areas of Algarve (the southern
province of Portugal), where the species ap-
pears to be particularly common, with the
exception of two specimens only that were
collected in Sesimbra (south of Lisbon). The
localities and sample sizes are C. orni: Albu-
ritel, Vila Nova de Ourem (n = 32); C. har-
hara hisitanica: Carvoeiro (n = 25), Praia da
Rocha (n = 4), Serra de Monchique (n = 1),
and Sesimbra (n = 2).

Methods

Measurements and Counts

Thirty-seven of the 40 characters were mea-
surements; the remaining 3 were counts.
Measurements were made using a Wild M3
microscope with a graduated eyepiece and
were taken as described in Table 2 or as illus-
trated in Figures 1-9. Of these 40 characters,
33 refer to external morphology and the re-
maining 7 to male genitalia.

Data Analysis

Data processing was carried out on the
CDC 6500 computer at the Imperial College
Computer Center (University of London) us-
ing two multivariate statistical programs de-
veloped by Prof. R. G. Davies (Department of
Pure and Applied Biology, Imperial College)
for cluster analysis and ordination (Quartan
and Davies 1983, 1985).

In most analyses, characters were standard-
ized by expressing each state as a deviation
from the mean in standard deviation units.

For Q-mode analysis, taxonomic distances
as well as product-moment correlations were
found and structured by the WPGMA method
of cluster analysis (Weighted Pair Group
Method with Arithmetic Averaging). Pheno-
grams, expressing the phenetic relationships
among the OTUs in a hierarchy of increasingly
larger clusters, were thus obtained (Figs.
10-16).

For R-mode analysis, character correlations

Table L List of specimens (males) oi Cicada orni Lin-
naeus and of C. harhara lusitanica Boulard investigated

(OTUs).

OTUs Locality, date of capture, and collector

Cicada orni

1-15 Alburitel, 10.viii.l979, J. A. Quartau
16-32 Alburitel, vii. 1971, J. A. Quartau

C. harhara hisitanica

33-34 Carvoeiro, 14.viii.l966, P. D. Rodrigues

35-38 Praia da Rocha, IT.viii. 1973, J. A. Quartau

39-40 Carvoeiro, 14. vii. 1978, J. A. Quartau

41-47 Carvoeiro, 30. vii. 1978, J. A, Quartau

48-55 Carvoeiro, 28. vii. 1978, J. A. Quartau

56-57 Carvoeiro, 9.viii-10.i.x.l980, L. Mendes

58-59 Sesimbra, 2.viii. 1980, J. A. Quartau

60 Carvoeiro, 31. vii. 1978, J. A. Quartau

61-62 Carvoeiro, 18. vii. 1978, J. A. Quartau

63 Monchique, 2.i.x.l971, F. Carvalho

64 Carvoeiro, 24.viii.1981, J. A. Quartau

based on data standardized by OTUs were
subjected to principal component analysis
(PGA). This ordination method transforms the
original characters, generally continuous, cor-
related characters, into a suite of uncorre-
cted, composite variables — the principal
components (principal axes). In addition to
being mutually independent, these compo-
nents account for maximum variance as fol-
lows: the variance along the first axis (i.e., the
corresponding eigenvalue) is the maximum
possible. The second axis describes the next
largest variance orthogonal to (uncorrected
with) the first. The third axis follows similarly
but is independent of both first two axes, and
so on, for as many axes as one wishes to extract
(e.g., Gibson et al. 1984). A transposed matrix
of the character loadings was post-multiplied
by the standardized data matrix to yield a
matrix of OTU projections in the principal
component space. Two-dimensional ordina-
tion diagrams of the representations of the two
species, together with the character loadings
(scaled eigenvectors), were thus obtained
(Figs. 17-18, Table 3).

Results

Phenograms

The seven phenograms resulting from vari-
ous analyses based on all characters, on the
genitalia only, or on the external characters
alone are shown in Figures 10-16. The
WPGMA clustering technique was followed

1988

QuARTAU: Cicada Analysis

173

Table 2. Description of characters: measurements
and counts (terminology mostly follows Myers [1928]).

Character No.

Description

1. Overall length measured from tip of crown to apical
margin of the right forewing with the latter in posi-
tion of rest alongside the body (Fig. 1).

2. Length of crown measured along a medial line pass-
ing through the median ocellus (Fig. I).

3. Minimum distance between the ocular sutures mea-
sured along the paired ocelli (Fig. 2).

4. Medial length of frons measured dorsally as indicated
(Fig. 2).

5. Medial length of pronotum measured dorsally as in-
dicated (Fig. 1).

6. Medial length of mesonotum measured dorsally from
anterior margin to posterior margin of cruciform ele-
vation or scutellum (Fig. 1).

7. Width of pronotum measured at the level of anterior
lateral margins (Fig. 1).

8. Width of pronotum measured at the level of postero-
lateral margins (Fig. 1).

9. Width of crown measured at the level of median
ocellus and as indicated (Fig. 2).

10. Inner distance between the paired ocelli (Fig. 2).

11. Distance between the right paired ocelli and the
right ocular suture as indicated (Fig. 2).

12. Distance between the base of the left antenna and the
left ocular suture as indicated (Fig. 3).

13. Inner distance between the base of antennae (Fig. 3).

14. Length of frons as illustrated (Fig. 3).

15. Length of clypeus as illustrated (Fig. 3).

16. Length of the exposed part of beak.

17. Length of dorsal margin of the left fore femur as
illustrated (Fig. 5).

18. Length of ventral margin of the left fore femur as
illustrated (Fig. 5).

19. Length of basal spine in ventral margin of left fore
femur as illustrated (Fig. 5).

20. Length of apical spine in ventral margin of left fore
femur as illustrated (Fig. 5).

21. Distance between tips of the apical and basal spines
in the ventral margin of the left fore femur as illus-
trated (Fig. 5).

22. Distance from anterior right corner to posterior left
corner of left operculum as illustrated (Fig. 6).

23. Distance from anterior left corner to posterior right
corner of left operculum as illustrated (Fig. 6).

24. Length of right forewing as illustrated (Fig. 1).

25. Greatest width of right forewing as illustrated (Fig.
4).

26. Length of subcostal cell ("gancho" cell of G6mez-
Menor 1957) in right forewing (Fig. 4).

27. Length of anterior margin of basal cell in right
forewing (Fig. 4).

28. Length of posterior margin of basal cell in right
forewing (Fig. 4).

29. Maximum width of basal cell in right forewing.

30. Minimum width of basal cell in right forewing.

31. Number of apical cells in right forewing.

32. Number of cells other than apicals of right forewing.

33. Number of spots in cross-veins of right forewing.

34. Length of pygofer in lateral view as indicated (Fig. 7).

35. Overall length of tenth abdominal segment as indi-
cated (Fig. 7).

36. Overall length of appendages of tenth abdominal
segment as indicated (Fig. 7).

37. Distance in basal curvature of shaft of aedeagus as
indicated (Fig. 9).

38. Width of shaft of aedeagus as indicated (Fig. 9).

39. Width of shaft of aedeagus in area of curvature as
illustrated (Fig. 9).

40. Medial length of eighth sternite or hypandrium (Fig.

in all; and Pearson's product-moment coeffi-
cient and the taxonomic distance coefficient
were used as measures of taxonomic proxim-
ity.

(a) Genital analyses

Figures 10 and 11 illustrate a correlation
and a distance phenogram, respectively, both
based on the seven standardized variables of
the male genitalia. Both analyses, notwith-
standing their being based on a small number
of variables, resulted in two main clusters,
one with C. orni and the other with C. bar-
bara lusitanica. However, in the former phe-
nogram, the cluster oi barbara lusitanica in-
cludes one specimen of orni (No. 13).

(b) External characters

These analyses resulted in the production
of the phenograms depicted in Figures 12 and
13. Both were based on standardized data, but
only the correlation coefficient succeeded in

giving an almost complete separation of the
two species of cicadas. In fact, OTUs were
grouped into two main clusters as in the geni-
tal analyses, but specimen No. 19 belonging
to C. orni appeared misplaced within C. bar-
bara lusitanica (Fig. 12). On the contrary, the
distance phenogram provided much less satis-
factory results than the previous analysis,
since each of the two major clusters incorpo-
rates elements of both species of cicadas (Fig.
13).

The phenograms of this group of analyses,
involving all 40 characters combined, are il-
lustrated in Figures 14-16.

Considering the phenograms based on
standardized data (Figs. 14, 16), it is clear that
the correlation phenogram gave a much bet-
ter distinction between the two species than
the distance analysis. In fact, the latter (Fig.
16) clustered six specimens of C. orni with C.

174

Great Basin Naturalist Memoirs

No. 12

1 10 'i
i '

1 1 9 ' 1

1 ' 1 : n 1 :

Figs 1-6. Diagrams of a male Cicada orni Linnaeus illustrating most of the measurements taken; 1, body, dorsal
view; 2, head, dorsal view; 3, head, anterior view; 4, right forewing, dorsal view; .5, left foreleg, mner view; 6, lett
operculum, ventral view.

1988

QuARTAU: Cicada Analysis

175

Figs. 7-9. As in Figures 1-6: 7, pygofer and tenth abdominal segment, ventral view; 8, eighth stemite or
hypandrium, ventral view; 9, aedeagus, lateral view.

barbara lusitanica. Even when the data were
unstandardized, correlations gave a good pic-
ture of the relationships between these two
species (Fig. 15).

Principal Component Analysis
This analysis involved all 40 characters and
was computed from a between-character cor-
relation matrix based on data standardized by
OTUs.

As in similar analyses carried out with
leafhoppers of the genus Batracomorphus
(Quartan 1983), slightly more than half (54%)
of the total variation in the study was ex-
plained by the first three axes.

The first component accounted for 38.90%
of the variation in the data and is interpreted
as a contrast between the lengths of the
pygofer, tenth abdominal segment, or ap-
pendages of the latter and the width of the
shaft of the aedeagus. It does not represent
overall size as commonly is the case, since
many of the characters (Table 3) are not posi-
tively correlated with it (e.g., Jolicoeur and
Mosimann 1960, Blackith and Reyment 1971,
Baker 1980, Gibson et al. 1984, Shea 1985).
In fact, it must represent both size and shape
as has been pointed out by several authors

(Mosimann 1970, Oxnard 1978, Humphries et
al. 1981). A complete separation of C. orm and
C. barbara lusitanica was given by the dis-
crimination afforded by this axis, which is
probably close to the orientation of the opti-
mum discriminant function. The characters
loading most heavily on this component
(Table 3) are therefore of considerable taxo-
nomic interest, since they are diagnostic for
this pair of species. The highest negative
scores, in decreasing order, were for charac-
ters numbered 35 (length of tenth abdominal
segment), 34 (length of pygofer), and 36
(length of appendages of tenth abdominal seg-
ment). The highest positive score was for
character numbered 38 (width of shaft of
aedeagus).

The second principal component accounted
for 8.51% of the total variation and was inter-
preted as a contrast between the number of
spots in cross-veins of the wings and the width
of the crown. It was most heavily loaded, neg-
atively and positively, on characters num-
bered 33 and 9, respectively.

The third principal component accounted
for 6.61% of the total variation and was inter-
preted as a factor resulting from the lengths of

176

Great Basin Naturalist Memoirs

No. 12

Table 3. Eigenvector matrix (character loadings) in a
principal component analysis of the matrix of correlations
among the 40 variables (data standardized by OTUs.)

Variables

Scaled

eigenvectors

III

0.296

-0.1.58

0.441

0.404

0.123

0. 155

0.531

0.557

-0.174

0.843

-0.160

0.126

0.492

0.300

-0.119

0.037

-0.096

0.280

0.567

0.280

-0.069

0.314

0.365

-0.131

0.336

0.633

-0.1,33

0.738

-0.237

-0.000

0.687

0.155

-0.200

0.859

-0.127

-0.142

0.313

0.423

0.414

0.117

0.422

0.306

-0.604

-0.196

-0.120

-0.769

0.104

-0.192

-0.316

0.338

0,524

-0.667

0.174

0.,375

0.565

0.2.52

-0.091

0.531

-0.066

0.111

-0.132

0.019

0.257

0.784

0.203

0.160

0.847

0.112

0.000

0.524

-0,340

0.060

0.683

-0.104

0.154

0.187

-0.207

-0.412

-0.334

0.126

-0,561

0. 133

0.117

-0.,549

-0.011

-0.207

0.471

0.129

-0.287

0.511

0.818

-0.490

-0.049

0.818

-0.479

-0.059

0.266

-0.745

-0.163

-0.952

-0.011

-0.040

-0.958

-0.153

-O.OOI

-0.944

-0.180

0.043

-0.869

-0.250

0.028

0.942

-0.065

-0.023

0.892

-0.249

0.024

-0.858

-0.125

-0.090

Latent roots

15.559

3.406

2.644

Percentage of

component 38.898

8.514

6.609

variation

cumulative 38.898

47.412

,54.021

the anterior and posterior margins of the basal
cell of the wings. It was most heavily loaded
(negative scores) on characters numbered 27
and 28.

Neither the second nor the third axis leads
to a separation of the two species of cicadas.
Figures 17 and 18 are two-dimensional views
of the relationships among specimens of both
species in the space determined by compo-
nent I combined with component II and by

component I with component III, respec-
tively. It is clear that these two plots gave a
good separation between C. orni and C. bar-
bara lusitanica. The plot combining axes II
and III did not, however, succeed in giving a
correct assignment of the two species and
therefore was not illustrated here.

Conclusions

Cluster analysis and principal component
analysis are two of the methods most com-
monly used for recognition of group structure
in numerical ta.xonomy (e.g., Quartau 1987).
This study suggests that the two general
methods are also very useful for discriminat-
ing between pairs of closely related species.
In fact, because of the uniformity of the gen-
eral clustering pattern that resulted from the
application of both methods, it is clear that
two distinct species exist, a fact also indicated
by behavioral data.

However, concerning cluster analysis, it is
worthwhile noting that the hierarchical struc-
ture within each of the two major groups of
OTUs differs a good deal from one particular
method to another. Moreover, it appeared
that correlations were more effective than tax-
onomic distances in describing relationships
between the two cicadas, a result in keeping
with Boyce (1964) or Cheetam (1968), for in-
stance. It is interesting to note, in this regard,
that such finding is in disagreement with a
similar study carried out with leallioppers
(Quartau and Davies 1983) or with results
based on other groups (e.g.. Smith 1972). Fi-
nally, the failure of the cluster analysis using
distances with nongenital characters shows
that the use of such techniques needs to be
undertaken with care and that it might be best
to use a consensus of several clustering tech-
niques when applying numerical methods to a
novel taxonomic situation.

Principal component analysis succeeded in
giving a good distinction of the two species
along axis I, in spite of its accounting for only
.38.90% of the total variation. This analysis also
showed that the main distinguishing charac-
ters between C. orni and C. barbara lusitan-
ica are the following: lengths of the pygofer,
the tenth abdominal segment, and the ap-
pendages of the latter (which are smaller in C.
barbara lusitanica. Figs. 19, 22), as well as
the width of the shaft of the aedeagus (which is

1988

QuARTAU; Cicada Analysis

177

-0.8

Correlations
Data standardized
Genital characters

D C. orni

• C. barbara lusitanica

Distances

Data standardized

Genital characters

DC. orni

• C. barbara lusitanica

Figs. 10-11. 10, correlation phenogram based on the seven genital characters with standardized data; 11, distance
phenogram based on the seven genital characters with standardized data.

thinner in orni. Figs. 20, 23). Moreover, de-
tailed examination of the male genitalia also
showed that the dorsal spine of the pygofer is
smaller in barbara lusitanica than in orni
(Figs. 19," 22).

Acknowledgments

1 thank Prof. R. G. Davies, Department of
Pure and Applied Biology, Imperial College,
for having made his computer programs avail-
able for my use as well as for his critical com-
ments on the manuscript. Prof. R. M. Ander-
son is also acknowledged for having granted

working facilities at the Imperial College
(University of London). 1 am also grateful to
my colleague Dr. M. L. Mathias for her help
with the preparation of most phenograms, and
to Miss A. R. Cruz for help with other illustra-
tions and the measurements. For enabling me
to publish this study in honor of Prof. P. Oman
(Oregon State University, USA), who has
been a source of encouragement since the
beginning of my taxonomic studies on
leafhoppers, I also acknowledge Dr. M. W.
Nielson (Brigham Young University, USA)
and Dr. H. D. Blocker (Kansas State Univer-
sity, USA).

178

Great Basin Naturalist Memoirs

No. 12

Correlations

Data standardized

Cxt. Morphology characters
D C. orni
• F. barbara lusitanica

Figs. 12-13. 12, correlation phenogram based on the 33 external morphological characters with standardized data;
13, distance phenogram based on the 33 external morphological characters with standardized data.

This paper was completed at Imperial Col-
lege (University of London) under an INIC
(Ministerio da Educagao, Lisbon) sabbatical
fellowship.

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1988

QUARTAU: CiCADA ANALYSIS

179

0.0

0.993

0.998

Correlat ions
Data standardized
All characters

DC. orni

o C. barbara lusitanica

T 31

Correlat ions

Data unstandardized

All ctiaracters
n £. orni
o C. barbara lusitanica

Figs. 14-15. 14, correlation phenogram based on all 40 combined characters with standardized data; 15, correlation
phenogram based on all 40 combined characters with unstandardized data.

Humphries, J M , F L Bookstein, B Chernoff, G R
Smith, R L Elder, and S G Poss 1981. Multi-
variate discrimination by shape in relation to size.
Systematic Zoology 30: 291-308.

JOLICOEUR, P , AND J E MosiMANN 1960. Size and shape
variation in the painted turtle. A principal compo-
nent analysis. Growth 24: 339-354.

MosiMANN, J E 1970. Size allometry: size and shape
variables with characterizations of the lognormal
and generalized gamma distributions. Journal of
the American Statistical Association 65: 930-945.

Myers, J G 1928. The morphology of the Cicadidae
(Homoptera). Proceedings of the Zoological Soci-
ety of London 1928: 265-472.

OXNARD, C E 1978. One biologist's view of morphomet-
ries. Annual Review of Ecology and Systematics 9:
219-241.

QuARTAU, J A 1983. A comparison of different numerical
methods applied to the taxonomy of Batracomor-
phus Lewis (Insecta, Homoptera, Cicadellidae,
lassinae): ordination techniques. Journal of the
Entomological Society of Southern Africa 46(1):
9-35.

1987. Ordination vs. cluster analysis in the numer-
ical taxonomy of some leafhoppers (Homoptera,
Cicadellidae). Pages 207-215 in M. R. Wilson and
L. R. Nault, eds.. Proceedings of the 2nd Interna-
tional Workshop on Leafhoppers and Planthop-
pers of Economic Importance, CLE., London.

QuARTAU, J A , AND R G Davies 1983. A comparison of
different numerical methods applied to the taxon-
omy of Batracomorphus Lewis (Insecta, Ho-
moptera, Cicadellidae, lassinae): cluster analysis.
Revista de Biologia 12: 550-596.

180

Great Basin Naturalist Memoirs

No. 12

Dis

tnnrcs

Dat

a slanda

diziMl

All

characters

DC.
or.

larbara

usil.iruc-.-l

•

; • ••-.=

•.

• . • •

•

orni

barbara lusicanica

D D D

* •

□ a

barbara lusiiamca

Figs. 16-18. 16, distance phenogram based on all 40 combined characters with standardized data; 17, a two-dimen-
sional view of the relationships among the 64 OTUs in a space determined by component I on ,x-axis (38.90%) and
component II on y-axis (8.51%) of a principal component analysis of the matri.x of correlations among all 40 characters
with standardized data; 18, as in Fig. 17 but referred to component I on x-axis and III on y-axis (6.61%).

1985. Character selection by information content

in the numerical taxonomy of some male Batraco-
morphus (Homoptera: Cicadellidae). Zeitschrift
fiir zoologische Systematik und Evolutionsfor-
schung 23; 100-115.

QUARTAU, J a . AND P. J. FONSECA. 1988. An annotated
check-list of the species of cicadas known to occur
in Portugal (Homoptera: Cicadoidea). Pages 367-
375 in C. Vidano and A. Arzone, eds. Proceedings
of the 6th Auchenorrhyncha Meeting, Turin.

Shea, B T 1985. Bivariate and multivariate growth
allometry: statistical and biological considerations.
Journal of Zoology 206: 367-390.

Smith, J D 1972. Systematics of the chiropteran family
Marmoopidae. University of Kansas, Publications
of the Museum of Natural History 56: 1-132.

Temple, J T 1968. The Lower Llandovery (Silurian) bra-
chiopods from Keisley, Westmorland. Palaeon-
tographical Society 521: 1-58.

1988

QuARTAU: Cicada Analysis

181

Figs. 19-24. Diagrams of the male genitalia oiCicada orni (Figs. 19-21) and of C. barbara lusitanica (Figs. 22-24):
19, 22, pygofer and tenth abdominal segment, lateral view; 20, 23, aedeagus, lateral view; 21, 24, eighth sternite or
hypandrium, ventral view (scale = 0.5 mm).

REVISION OF THE NIRVANINAE (HOMOPTERA: CICADELLIDAE)
OF THE INDIAN SUBCONTINENT

C. A. Viraktamath' andC. S. Wesley^

Abstract — ^Three tribes of Nirvaninae, namely, Balbillini (two genera and three species), Nirvanini (five genera and
26 species), and Occinirvanini (one genus and one species), occur on the Indian subcontinent. The tribes, genera, and
species are redescribed, illustrated, and keyed. New ta.xa recognized are BalbiUus indicus n. sp. (India; Kerala),
Stenotortor stibhimalaya n. sp. (India: West Bengal), Kana bispinosa n. sp. (India: Tamil Nadu), K. nigropicta n. sp.
(India: Kerala), Sophonia bakeri n. sp. (India: Karnataka, Kerala, Tamil Nadu, Uttar Pradesh; Nepal), S. bifida n. sp.
(India: West Bengal), S. complexa n. sp. (India; Meghalaya), S. compUcata n. sp. (India: Mizoram, Meghalaya), S.
keralica n. sp. (India; Kerala), Nirvana peculiarisn. sp. (India: Mizoram, Meghalaya), and N. striata n. sp. (India; West
Bengal, Uttar Pradesh, Himachal Pradesh). The genus Qitercinirvana Ahmed & Mahmood (type species; Q. longi-
cephala Ahmed & Mahmood) is treated as a junior synonym oi Sophonia Walker. Kana signata Distant and Nirvana
greeni Distant are suppressed as junior synonyms of K. illuminata and N. Hnealis , respectively, and are transferred to
Sophonia . Three other species transferred to Sophonia are Kana modesta Distant, Quercinirvana longicephala Ahmed
& Mahood, and Nirvana insignis Distant. The relationships among various genera are discussed briefly. The genera
Crispina Distant and Mohiinia Distant are excluded from Nirvaninae.

The Nirvaninae, one of the smaller sub-
families of the leafhopper family Cicadellidae,
include a predominantly tropical group of
leafhoppers. They are fragile, often brightly
colored, depressed leaflioppers and common
on lush green vegetation. They are often mis-
taken for Typhlocybinae, but the structure of
the head and the more depressed form and
truncate basitarsus of the hind leg distinguish
Nirvaninae. The subfamily includes 111 spe-
cies (excluding Mukariinae and other non-
Nirvaninae genera, which were traditionally
included in the Nirvaninae) distributed in the
Oriental (56), Afrotropical (24), Neotropical
(7), Palearctic (7), Australian (11), and Pacific
island (6) regions. The fauna of Afrotropical
(Linnavuori 1979), Australian (Evans 1966),
Papua New Guinea (Evans 1973), Neotropical
(Kramer 1964), and Palearctic (Kuoh and
Kuoh 1983) regions have recently been re-
vised. The tribe Mukariini earlier included in
the subfamily by Distant (1908g) and Metcalf
(1963) is now considered a subfamily distinct
from Nirvaninae (Linnavuori 1979).

Information on host plants of Nirvaninae is
very meagre. Nirvana pallida Melichar and
N. suturalis Melichar are destructive to sugar-
cane and grasses in Taiwan (Schumacher
1915a). They also feed on rice, mulberry, and
camphor tree (Esaki and Ito 1954a). Baker

(1923a) found Stenotortor inocarpi Baker on
Otaheite chestnut, Inocarpus edulis, in Sin-
gapore. In Australia, Nirvaninae feed on /.
edulis, casuarina, and croton (Evans 19411).
Teco7naria capensis is recorded as the host of
Narecho tecomariae Theron in South Africa
(Theron 1970). Quercinirvana longicephala
Ahmed & Mahmood feeds on Aesculus indica,
Quercus dilatata, and Vihurmmi nervosum
(Ahmed and Mahmood 1970). Chudania
delect a Distant breeds on Ficus carica
(Ahmed and Mallik 1972). In South America,
Tahura fowleri Kramer is found on Passiflora
(Kramer 1964). Kuoh and Kuoh (1983)
recorded Camellia sinensis. Acacia confusa.
Citrus reticulata, Pterocarpus indicus, Orijza
sativa, Psidium guajava, and Chimonanthus
praecox as host plants of species oi Pseudonir-
vana Baker { = Sophonia Walker) in China.
Nirvana pallida and N. greeni Distant breed
on black gram, green gram, cowpea, field
bean, pigeon pea, and soybean in India (Ra-
makrishna 1980).

The Nirvaninae of the Indian subcontinent
consist of eight genera and 30 species grouped
in three tribes, namely, Balbillini, Nirvanini,
and Occinirvanini. The genera Mohunia Dis-
tant and Crispina Distant are excluded from
the study, as they probably belong to the sub-
family Deltocephalinae.

Department of Entomology, University of Agricultnral Sciences. CKVK Campus, Bangalore 560 065, India.
^Department of Ecology and Evolution, S.U.N.Y., Stony Brook, New York 11794, US.\.

182

1988

VlRAKTAMATII, WESLEY: REVISION OF INDIAN NiKVANINAE

183

Material and Abbreviations

This study was based mainly on specimens
collected over the years and deposited in the
insect collection of the Department of Ento-
mology, University of Agricultural Sciences
by the senior author and his students. Addi-
tional specimens were borrowed from various
institutions for the study. Abbreviations used
for these institutions and for the depositories
of the types of new taxa are as follows:

BMNH - British Museum (Natural History), London.

lARI -Indian Agricultural Re.search Institute, New
Delhi.

IRSNB - Institute Royal des Sciences, Naturelles de Bel-
gique, Bru.xelles.

JU - Department of Biosciences, Jammu Univer-

sity, Jammu.

PAU - Punjab Agricultural University, Ludhiana.

UAS - University of Agricultural Sciences, Bangalore.

USNM -U.S. National Museum (Natural History),
Washington, D.C.

ZMHU -Museum fiir Naturkunde der Humboldt, Urii-
versitat zu Berlin, Berlin.

ZSI -Zoological Survey of India, Calcutta.

Distant (1908g, 1918b) did not mention the
number of specimens (syntypes) of each new
species he described, except when a single
specimen was involved, viz., Chundania de-
lecta Distant (1908g; 268) In this study lecto-
type designations were made even though a
single specimen of the type series (syntypes?)
was present in the BMNH collection. Unless
otherwise stated they were considered as part
of the syntype series.

The terminology used for describing hind
leg spinulation follows Davis (1975).

Description of Taxa

Nirvaninae

Delicate, fragile, depressed, small to mod-
erately large (4.0-9.0 mm) leafhoppers. Usu-
ally yellow or white, often marked with red,
black, orange, or brown fascia. Head as wide
as or wider than pronotum. Vertex usually
flattened, strongly produced, and with rugae
or sculpturing on lateral and anterior regions.
Ocelli on crown along lateral margin in front of
eyes. Frontoclypeus and clypellus either flat
or tumid, in the latter case often provided
with a median keel. Lora often small, extreme
anterior margin of genae bluntly prominent,
extending beyond apex of clypellus. Antennal
pits deep, antennal ledges more or less
strong, antennae long. Lower margin of eyes

sinuate ventrally. Anterior tentorial branches
L-shaped. Lateral margins of pronotum cari-
nate (except in Omaranus Distant), moder-
ately long. Forewing venation reduced, with-
out cross-veins at base (except Balbillini),
longitudinal veins represented by a series of
paired pits basally, appendix either narrow
(Nirvanini) or broad (BcdhiUus Distant). Hind-
wing with three or lour apical cells, veins lA
and 2A fused basally. Fore and middle tibiae
cylindrical or flattened (Balbillini), hind
femoral spinulation 2+1 + 1 (Nirvanini),
2+1 + (Balbillini and Occinirvanini), or
2+0+0 (Balbilhni).

Male pygofer either cylindrical or de-
pressed, with or without anal collar process,
but may be armed with ventral process, heav-
ily macrosetose. Valve small, fused with
pygofer laterally. Plates usually parallel-
sided. Style variable. Connective Y-shaped
(Nirvanini) or platelike (Balbillini). Aedeagus
with single shaft and often provided with pro-
cesses.

Linnavuori (1979) suggested that the sub-
family Nirvaninae is a derivative of the Aphro-
dinae-Cicadellinae stock and that Nirvanini is
the most advanced tribe.

Key to Tribes

1. Fore and middle tibiae flattened; head notched
in front of eyes so that the scape is visible in
dorsal aspect Balbillini

— Fore and middle tibiae rounded; scape not visi-
ble from above 2

2(1). Ocelli nearer to apex of head than to eyes;
forewing appendix wide and extending around
apical wing margin; lora large; vertex about half
as long as wide, obliquely produced in front of
eyes, with hind femoral spinulation 2+1 + 0;
hind basitarsus with six platellae . . . Occinirvanini

— Ocelli nearer to eyes than to apex of head;
forewing appendix narrow and does not extend
around apex of wing; lora small; vertex as long as
wide or longer, not obliquely produced in front
of eyes; hind femoral spinulation 2+1 + 1; hind
basitarsus with two or three platellae . . Nirvanini

Tribe Balbillini

This tribe has been well characterized by
Linnavuori (1979). The following additional
characters are noted. Forewing with supernu-
merary cross-veins along costal margin and
with two m-cu cross-veins, four to five apical
cells, two anteapical cells, and appendix (ab-
sent in Stenotortor Baker). Hindwing with
four apical cells. Hind femoral spinulation

184

Great Basin Naturalist Memoirs

No. 12

2 + 2 + or 2 + 0+0. Male connective long and
incrassate or lamellate and fusiform. Aedea-
gus simple with large gonopore.

Key to Genera of Balbillini

1. Forewing with a well-developed appendix; hind
femoral spinulation 2+1+0 Balhillus Distant

— Forewing without an appendix; hind femoral spin-
ulation 2 + + Stenotortor Baker

Genus Balhillus Distant

Balhillus Distant 1908g: 287. Type species; Balhillus
granulosus Distant, by original designation and
monotypy.

Head narrower than pronotum. Crown
bluntly, angularly produced, convex, de-
clivous anteriorly, with median and submar-
ginal, anterior carinae between ocelli. Face
horizontal, flat. Clypellus twice as wide at
base as at apex; lora small. Lateral margins of
head in front of eyes notched, exposing scape
of antenna. Pronotum declivous laterally,
with two lateral, carinate margins; mese-
pisternum attaining lateral position in front of
base of wing and strongly ridged. Fore and
middle tibiae angular; hind femoral spinula-
tion 2+1 + 0, inner spine very slender and
minute. Apex of hind basitarsus with three
platellae. Hind tibial spinulation Rj 12 ± 1, R,
19±1, R3 25±l, R4 25±l.

Pygofer much shorter than its height, with-
out processes. Anal tube short. Genital plate
parallel-sided, 3.4 times as long as wide with
uniseriate, submarginal setae and few hairlike
setae. Apophysis of style simple, without
preapical lobe. Connective lamellate and fusi-
form. Aedeagus articulate with connective,
simple and with large gonopore.

Remarks. — As pointed out bv Linnavuori
(1979) and Baker (1923), Balhillus and Steno-
tortor are closely related. They can be sepa-
rated by the characters used in the key. The
bodies of both genera remain appressed to the
leaf surface, which explains the flat nature of
the tibiae and the undersurface of the body.

The Indian species, B. granulosa and B.
indica, have hind femoral spinulation 2+1 +
and three platellae at the apex of the hind
basitarsus. The male plate of B. indica is paral-
lel-sided. The connective and aedeagus are
articulated rather than fused. The connective
is spindle-shaped, lamellate rather than long
and incrassate, and the preapical lobe of the
style is wanting. These characters strongly

suggest that the Afrotropical species of Balhil-
lus, namely, B. ahas Linnavuori and B. tri-
maculatus Linnavuori, may not belong to Bal-
hillus.

Key to Species of Balhillus

1. Vertex and scutellum with dark brown spots ....
granulosus Distant

— Vertex and scutellum immaculate . . . indicus, n. sp.

Balhillus granulosus Distant
Figs. 1-5

Balhillus granulosus Distant 1908g: 288. Holotype 9 , Sri
Lanka (BMNH, examined).

Yellow. Vertex with a median spot, a spot in
apical half on either side divided by carina,
and a spot near inner margin of eye dark
brown. Pronotum with a median, short stripe
on anterior margin fuscous. Basal triangles on
scutum and apex of scutellum dark brown;
blackish, median line on scutellum not reach-
ing apex of scutellum. Longitudinal veins and
claval veins lemon yellow, entire surface of
forewing punctate, punctations along radial
vein and on outer apical cell and a spot at base
of appendix fuscous.

Vertex bluntly conical, 1.24 times as wide
between eyes as its median length, median
carina prominent in its apical half, apical 0.75
of disc sculptured, basal 0.25 polished. Face
about as long as wide. Pronotum widened
posteriorly. Scutellum longer than pronotum,
transversely, finely rugulose beyond sulcus.
Forewing 3.4 times as long as wide, with one
m-cu cross-vein abut 0.33 distance from base
and another near apex limiting the short, in-
ner anteapical cell, outer anteapical cell open
behind, numerous veins arising from apical
half of outer anteapical cell and reaching
costa, few of them forked at costal margin,
third apical cell divided by cross-vein. Hind-
wing with four apical cells.

Female genitalia. — Seventh sternum
about twice as long as sixth, its hind margin
slightly concave with a strong, U-shaped, me-
dian excavation, very finely, transversely
rugulose. Ovipositor exceeding pygofers.

Measurements. — Female 7.10 mm long,
head 1.85 mm wide, pronotum 1.87 mm
wide.

Spegimen e.xamined. — Holotype 9 la-
beled /Type/ /Balhillus granulosus Dist.,
tvpe/ /Ceylon (Green)/ /Distant Coll.,
1911-383/ (BMNH).

1988

ViRAKTAMATH, WeSLEY: REVISION OF INDIAN NiRVANINAE

185

/6 17

Figs. 1-17. Species oi Balbillus. B. granulosus Distant; 1, head and thorax; 2, same, profile; 3, face and part of
prothorax;4, forewing; 5, ovipositor. B. indicus, n. sp.: 6, general habitus; 7, head and thorax; 8, face; 9, head and
thorax, profile; 10, forewing; 11, pygofer; 12, male plate; 13, connective; 14, style; 15, 16, 17, aedeagus, lateral,
cephalic, and caudal views.

Remarks. — This species is closely related
to B. indicus, from which it differs by its
distinctive coloration.

Balbillus indicus, n. sp.

Figs. 6-17

Uniformly pale yellow. Apex of clavus.

186

Great Basin Naturalist Memoirs

No. 12

second and third apical cells of forewing suf-
fused with brown.

Disc of vertex polished, medially, longitu-
dinally grooved for 0.66 distance and ridged
beyond. Forewing 2.5 times as long as wide,
punctate along veins of corium, in cells of
clavus, and along the costal margin, m-cu
cross-veins two (as in B. granulosus) but not
easily seen, fewer oblique veins reaching
costal margin from outer anteapical cell.

Male genitalia. — Pygofer simple, three
times as long as its height with a few stout
setae on caudoventral area, anal tube short.
Plate parallel-sided, with single row of sub-
marginal, stout, short setae, and hairlike setae
along outer margin. Style without preapical
lobe, apophysis long, its apex pointed and
directed ventrad. Aedeagus with preatrium
elongate, dorsal apodeme unpaired, short,
shaft tubular for greater distance caudad, then
flattened and deflected anteriorly, apex
pointed; gonopore apical, large.

Measurements. — Male 5.50 mm long,
head 1.28 mm wide, pronotum 1.53 mm
wide.

Specimen examined. — Holotype S, India:
Kerala: Thekkadi, 27.iii.1977, s' Viraktamath
CoU. (UAS).

Remarks. — In coloration and size it is simi-
lar to B. alhellus Baker from the Philippines,
but differs in forewing venation in having five
apical cells rather than four as in the latter
species.

Genus Stenotortor Baker

Stenotortor Baker 1923a: 375, 377. Type species: Steno-
tortor inocarpi Baker, by original designation and
monotypy.

Stucture similar to Balhillus but differs in
the following respects. Orange with brick red
or reddish brown markings. Median carina of
vertex more prominent. Forewing without
appendix, with one m-cu cross-vein, both an-
teapical cells open behind. Hind femoral spin-
ulation 2+0+0. Hind tibial spinulation Rj 20,
R2 13, R3 10. Apex of hind basitarsus with
three platellae.

Male pygofer simple, its caudoventral area
thickly setose. Anal tube short. Male plate
parallel-sided without macrosetae or long,
hairlike setae. Style with well-developed,
preapical lobe. Connective triangular, articu-
lated with simple aedeagus. Gonopore large
on ventral margin.

Remarks. — Baker (1923a) differentiated
Stenotortor from Balhillus by its strongly de-
pressed body, tectiform tegmina with ex-
panded costal area, strongly curved outer
margin, and obscure venation. However,
more important differences in Stenotortor ap-
pear to be the absence of an appendix on the
forewing, the presence of one m-cu cross-
vein, and hind femoral spinulation that is
2+0+0.

Stenotortor subhiinalaija, n. sp.

Figs. 18-27

Orange yellow. Vertex with oblique band
on either side of median line, oblique line
laterally; pronotum with lateral, oblique
band, transverse band on scutellum dark red-
dish brown. Forewing with submarginal band
on costal area, a stripe on clavus along com-
missure at basal 0.33 and then obliquely ex-
tending to inner claval margin, a median
stripe connecting both claval stripe basally
and inward extension of submarginal stripe
both medially and caudally, dark reddish
brown.

Vertex bluntly conical, slightly longer than
its width between eyes. Face about as wide as
long. Clypellus strongly narrowed apically.
Lateral margin of pronotum strongly diverg-
ing, 2.7 times as wide as its median length.
Forewing 2.4 times as long as its width. Hind-
wing with four apical cells.

Male genitalia. — Pygofer twice as long as
height, caudoventral margin with short, stout
setae. Anal tube short. Male plate parallel-
sided, about four times as long as wide, caudal
apex rounded. Style with well-developed,
stout, preapical lobe, apophysis fingerlike.
Connective triangular. Aedeagus with elon-
gate preatrium, without dorsal apodeme,
shaft tubular in basal 0.33, then spatulate with
large gonopore on its ventral margin occupy-
ing 0.66 of its length.

Measurements. — Male 4.90 mm long;
head 1.18 mm wide, pronotum 1.55 mm
wide.

Specimens e.xamined. — Holotvpe 6, In-
dia: West Bengal: Sukna, 3.xi.l981, C. A. Vi-
raktamath Coll. no. 277 (UAS).

Remarks. — This species differs from S. in-
ocarpi in coloration of the vertex, pronotum,
and wing pattern, in its larger size, and in its
relatively longer aedeagus.

1988

ViRAKTAMATH, WeSLEY: REVISION OF INDIAN NiRVANINAE

187

Figs. 18-27. Stenotortor stibhimalaya, n. sp.: 18, habitus; 19, face; 20, head and thorax, profile; 21, forewing; 22,
pygofer; 23, male plate; 24, connective; 25,26, aedeagus, lateral and caudal views; 27, style.

Tribe Nirvanini

Pale yellow with dark or red markings.
Body long, narrow, more or less depressed.
Head as long as or longer than its width, disc
of vertex longitudinally rugose in anterior re-
gion. Ocelli near lateral margins in front of
eyes. Face with anterior, median carina and
usually with lateral, oblique ridges on fronto-
clypeus. Gena narrow, lora small. Antennal
pits rather deep, near anterior margin of eyes
not visible from above; antennae long, reach-
ing at least hind margin of pronotum.
Forewings narrow, with reduced appendix
and venation indistinct basally, second api-
cal cell not widened apically. Hindwing
with three or four apical cells. Fore and mid-

dle tibiae rounded. Hind femoral spinulation
2+1 + 1. Hind basitarsus terminated by two
platellae.

Pygofer with well-developed, lateral lobes.
Anal tube elongate, often with apical or cau-
doventral process. Male plate elongate, Ion-
longer than pygofer and apically rounded.
Style variable, usually with slender apophysis
that is curved, hooked, or avicephaliform.
Connective free, Y-shaped. Aedeagus weakly
sclerotized, usually with appendages.

Remarks. — This group is most abundant
among Nirvaninae and appears highly
evolved compared to other tribes, consider-
ing the reduced wing venation, the more
streamlined body form, and the specialized
male genitalia.

188

Great Basin Naturalist Memoirs

No. 12

Key to Genera of Nirvanini
1. Hindwing with four apical cells (Fig. 59) 2

— Hindwing with three apical cells (Fig. 100) .... 3

2(1). Most parts of head, pronotum, and scutellum
black dorsally; frontoclypeus and clypellus tu-
mid (Figs. 85, 86); male connective rather T-
shaped (Fig. 95) Chudania Distant

— Dorsum of head, pronotum, and scutellum at
most with black, narrow stripe terminated by
round spot (Figs. 50, 60); frontoclypeus and
clypellus flat (Figs. 29, 45, 49); male connective
Y-shaped (Figs. 32, 75) Kana Distant

3(1). Crown of head depressed in middle, margined
by carina around its margin (Figs. 268, 269);
male plate with angulate projection on its lateral
margin near apex (Fig. 272); male style short,
stout, appearing like loosely closed fist (Fig. 273)
Ophiuchus Distant

— Crown of head either flat or convex; male plate
either smoothly rounded near apex or with
spine; style variable (Figs. Ill, 148, 160, 182),
slender and with slender apophysis that is
curved, hooked, or avicephaliform 4

4(3). Dorsal apodeme of aedeagus robust, bearing
processes at least at base of shaft (Figs. 118, 139,
150, 171, 180, 206); male plate about four times
as long as its width at its midlength (Figs. 114,
149, 170, 196); frontoclypeus with prominent,
lateral ridges Sophonia Walker

— Dorsal apodeme of aedeagus slender, U-
shaped, without prominent processes (Figs.
225, 240, 254, 267); male plate five times or
more as long as its width at its midlength (Figs.
227, 264); frontoclypeus with weak, lateral
ridges Nirvana Kirkaldy

Genus Kana Distant

Kana Distant 1908g: 285. Type species: Kana thoracica
Distant, by original designation.

Pale yellow, often with bright-colored
patches and stripes. Crown slightly (less than
1.5 times) longer than its width. Ocelli placed
laterad of submarginal carina. Frontoclypeus
with anterior ridge, rather flattened, with lat-
eral, very prominent, ridgelike folds. Clypel-
lus large, broad at base. Lora small. Face
slightly longer than wide between eyes. Sec-
ond apical cell of forewing parallel-sided.
Hindwing with four apical cells. Hind tibial
spinulation R^ 20, R. 12 ± 1, R, 23. Hind ba-
sitarsus with two platellae.

Pygofer rounded caudally, with one or two
rows of submarginal macrosetae; ventral pro-
cess robust at base, narrowed apically. Male
plate elongate, narrow, with outer margin sin-
uate. Connective Y-shaped, with arms joining
broadly; apophysis robust, short, with slender
apical extension. Aedeagus partially sclero-

tized, often depressed, with lateral, lamellate
margin, gonopore surrounded by membra-
nous tube with or without process, dorsal
apodeme reduced.

Remarks. — Some species of Kana, espe-
cially K. bispinosa and K. nigropicta, resem-
ble externally the species of Sophonia. The
four closed apical cells in the hindwing, how-
ever, distinguish this genus from Sophonia
and Nirvana. The aedeagus of Kana is charac-
teristic in that it is bent in its apical half and
bears appendages surrounding the gonopore;
the second apical cell on the forewing is not
narrowed apically as is the case in both Sopho-
nia and Nirvana. Kana is closely related to
Yaoundea Linnavuori and even may prove to
be synonymous with it. Both genera are re-
lated to Chudania, from which they differ in
having a flatter frontoclypeus.

Key to Species of Kana
1. Vertex, pronotum, and scutellum traversed by a
median, longitudinal, black line (Figs. 50, 60);
third apical cell of forewing with large, round,
black spot (Figs. 58, 61) 2

— Vertex or pronotum or both either immaculate
or with yellow, reddish, or orange stripes (Figs.
42, 44); forewing with brown or reddish orange,
oblique, or transverse bands, black spot on third
apical cell, if present verv small (Figs. 36, 45, 47,
79) ; 3

2(1). Vertex without black spot near apex (Fig. 50);
aedeagal shaft with pair of pronglike processes

(Fig. 55) bispinosa, n. sp.

— Vertex with black spot near apex (Fig. 60);
aedeagal shaft without processes (Fig. 67) ....
nigropicta, n. sp.

3(1). Forewing with transverse bands onlv (Figs. 36,

79) ' 4

— Forewing with both oblique and transverse
bands (Figs. 43, 47) 7

4(3). Forewing with single, broad, mottled brown,
transverse band, area beyond apical cross-vein
fuscous; vertex and pronotum marked with
white stripes; male genitalia as in Figs. 74-78
illahorata Distant

— Forewing with two bands either both yellowish,
orange, or red, or one orange red (more basal)
and the other fuscous (median); apical cells at
most with a few fuscous spots 5

5(4). Pronotum without colored stripes; vertex with
clear yellow, longitudinal stripe on each lateral
side; forewing with two transverse, yellow fas-
ciae decora (Melichar)

— Pronotiun with orange stripes; vertex either im-
maculate or with orange-yellow, submarginal,
apically converging stripes; forewing with basal,
orange-yellow or red, and median, fuscous,
transverse stripe 6

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189

Figs. 28-36. Kana ordinate Distant; 28, head and thorax; 29, same, profile; 30, ventral pygofer process; 31, male
plate; 32, connective; 33, style; 34, connective and aedeagus, lateral view; 35, aedeagus, caudal view; 36, forewing.

6(5). Vertex with pair of anteriorly converging, or-
ange-yellow stripes continued on pronotum . . .
fasciata Pruthi

— Vertex without stripes; pronotum with four or-
ange stripes; male genitalia as in Figs. 30-35 . .
ordinata Distant

7(3). Pronotum with two median, red stripes and two
lateral, anterior, lemon yellow spots (Figs. 44);
forewing with two red, transverse bands on
corium (Fig. 47) thoracica Distant

— Pronotum with anterior margin and two spots on
posterior margin orange-yellow (Fig. 42);
corium of forewing with three transverse, red
bands (Fig. 43); male genitalia as in Figs. 37-41
ramificata Distant

Kana ordinata Distant

Figs. 28-36

Kana ordinata Distant 1908g: 287. Lectotype 6, Sri
Lanka (BMNH, examined).

Yellow. Pronotum with two inner and two
outer orange stripes joined posteriorly in
male. Scutellum with brownish basal trian-
gles. Forewing with reddish orange band
across basal area, fuscous speckled band about
midlength and series of fuscous spots at apices
of anteapical cell.

Head slightly narrower than pronotum.
Vertex of head slightly longer than its width
between eyes (112:123), apex bluntly conical.
Face with four prominent, lateral ridges.

Male genitalia. — Pygofer rather squarish
with caudodorsal area angulate, and with two
rows of stout, long setae; ventral process with
triangular, dorsal lobe and median, dorsal
hump. Plate with rounded, caudal apex, with
row of stout setae in caudal 0.66 and with

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marginal row of long, hairlike setae. Style
with well-developed, preapical lobe, apophy-
sis short, stout, its apical extension slender,
avicephaliform. Connective Y-shaped with
stem as long as the length of arms. Aedeagus
tubular with unsclerotized, poorly developed
dorsal apodeme. Shaft strongly bent and di-
rected ventrad and cephalad in apical 0.33,
appearing rather like a compressed, inverted
J, cephalic part of shaft laterally expanded and
constricted in middle in cephalic view; caudal
part with two long, curved processes that
cross over and are longer than those in K.
ramificata.

Measurements. — Male body 3.00 mm
long, forewing 4.10 mm long, head 1.17 mm
wide, pronotum 1.25 mm wide. Female 6.40
mm long, head 1.30 mm wide, pronotum 1.32
mm wide.

Specimens examined. — Lectotype S la-
beled /Bogawantalawa, Ceylon, April/ /Dis-
tant Coll., 1911-383/ /1625/ /Nirvana ordi-
nata Dist., type/ here designated (BMNH).
Paralectotype 9 labeled /Maskeliya, Ceylon,
8-05/ /Distant Coll., 1911-383/ here desig-
nated (BMNH).

Remarks. — Kana ordinata is closely re-
lated to K. ramificata and differs in coloration
and structure of the pygofer process. Kana
ordinata has a shorter caudal part of the
aedeagus and longer processes that cross
over.

Kana ramificata Distant

Figs. 37-43

Kana ramificata Distant 1908g: 286. Lectotype 6, Sri
Lanka (BMNH, examined).

Pale yellow, with submarginal band on ver-
tex, anterior margin of pronotum and two
spots on posterior margin orange. Scutellum
golden yellow. Forewing with basal band con-
nected by similar but more oblique band from
costa to clavus and then continued on clavus
but not reaching claval tip, orange; another
oblique band with well-defined, orange, ante-
rior margin and cross-veins limiting apical
cells, reddish; spot at apex of clavus and an-
other on third apical cell fuscous.

Head slightly narrower than pronotum
(46:49). Vertex slightly longer than width be-
tween eyes, bluntly conical. Lateral ridges on
face four in number, not prominent. Scutel-
lum longer than pronotum.

Male genitalia. — Pygofer rather squar-
ish, with caudal tufts of long, stout setae; ven-
tral process stout with dorsal, triangular pro-
cess near caudal apex. Anal tube stout. Male
plate with oblique row of stout setae in caudal
0.66, with long, hairlike setae, apex bluntly
rounded. Style and connective as in K. ordi-
nata. Aedeagus as in K. ordinata, but caudal
bent part 0.66 as long as tubular cephalic part,
apical processes shorter.

Measurements. — Male 5.20 mm long,
head 1.15 mm wide, pronotum 1.22 mm
wide.

Specimen examined. — Lectotype S la-
beled/Green, Ceylon, 95-221/ /Ntroana ram-
ificata Dist., type/ here designated (BMNH).

Remarks. — Kana ordinata and K. ramifi-
cata share a similar basic pattern of male geni-
talia; however, they differ in the relative size
of the aedeagal processes and the shape of the
pygofer process. Both hindwings and the right
side of the forewing of the lectotype are dam-
aged and were placed in a microvial.

Kana thoracica Distant
Figs. 44-48

Kana thoracica Distant 1908g; 285. Lectotype 9 , Sri
Lanka (BMNH, examined).

Pale yellow. Two anteriorly converging,
submarginal stripes on vertex, two stripes one
on either side of median line red, two lateral
spots on anterior margin of pronotum lemon
yellow. Forewing with red bands (Fig. 47)
along anal margin, obliquely crossing over to
clavus; short band on corium near base, an-
other longer, oblique band about midlength
from costal margin to clavus. Series of spots on
either side of cross-veins separating apical
cells fuscous.

Head narrower than pronotum (46:50).
Vertex longer than its width between eyes
(37:26). Face with three lateral, oblique
ridges; longer than width including eyes.
Pronotum shorter than both vertex (24:37)
and scutellum (24:28).

Female genitalia. — Hind margin of sev-
enth sternum slightly concave with a median
protuberance.

Measurements. — Female 5.90 mm long,
head 1.15 mm wide, pronotum 1.25 mm
wide.

Specimen examined. — Lectotype 9 la-
beled /Pundaloya, Ceylon/ /Ceylon, Green

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191

Figs. 37-43. Kana ramificata Distant: 37, male pygofer; 38, connective and aedeagus, lateral view; 39, same, caudal
view; 40, male plate; 41, style; 42, head and thorax; 43, forewing.

Coll. 90-115/ /Nirvana thoracica Dist., type/
here designated (BMNH).

Remarks — This species appears to be re-
lated to K. ramificata and K. ordinata, from
which it differs in coloration.

Kana bispinosa, n. sp.
Figs. 49-59

Pale yellow. Vertex with median, longi-
tudinal, black line from before apex and ex-
tending to the tip of scutellum, gradually
widening posteriorly, lateral margin orange.
Pronotum reddish. Forewings with suffused,
red stripe adjoining thin, light brown stripe
terminating on claval suture at 0.66 distance
from base on anal margin, corium pinkish
distally, claval apex and apical wing mar-
gin brown, third apical cell with large, black
spot, surrounding area light brown, costal

margin with three reddish fascia.

Head narrower than pronotum. Vertex
slightly longer than width between eyes, flat,
slightly depressed about center, apex bluntly
conical, granulose. Apical half of face con vex,
basal half depressed, with three oblique
ridges.

Male genitalia. — Pygofer rounded, with
stout, long setae along caudal margin; ventral
process broad basally, abruptly narrowed and
curved dorsally near apex. Male plate about
four times as long as wide, distal 0.66 with an
oblique row of stout spines and with marginal,
hairlike setae. Style with stout, preapical lobe
and short, stout apophysis, apical extension
slender, beaklike. Connective Y-shaped with
arms as long as stem. Aedeagus with short,
well-developed preatrium; shaft with ridged

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Figs. 44-48. Kana thoracica Distant: 44, head and thorax; 45, head, profile; 46, face; 47, forewing; 48, female
seventh sternum.

lateral margins, somewhat rectangular, me-
dian ridge on caudal aspect running from base
to apex, gonopore surrounded by membra-
nous, tubelike structure bearing pair of ven-
trally directed, pronglike processes.

Measurements. — Male 4.50 (4.50-4.60)
mm long, head 0.93(0.90-0.95) mm wide,
pronotum 0.99 (0.98-1.00) mm wide.

Specimens examined. — Holotype d, In-
dia: Tamil Nadu: Naduvattam, 6.vi.l977, C.
A. Viraktamath Coll. (UAS). Paratypes 2 6,
data as in holotype but collected by S. Virakta-
math (BMNH, UAS).

Remarks. — This species is closely related
to Kana nigropicta but only distantly related
to K. illaborata and K. ordinata. It differs
from the latter two species in coloration of the
head, thorax, and forewing and in the struc-
ture of male genitalia.

Kana nigropicta, n. sp.
Figs. 60-68

Uniformly orange-yellow. Vertex with
round, black spot near apex, contiguous with
median, longitudinal, black line extending to
tip of scutellum. Lateral margins of vertex and

pronotum orange. Forewing with irregular,
hyaline patches, claval apex brown, third api-
cal cell with large, black spot, apical margin
reddish, costal margin with three red, oblique
fasciae, black spot in fourth apical cell.

Head as wide as pronotum. Vertex about
1.5 times as long as wide, anterior half convex,
faintly granulose. Apical 0.25 of face slightly
bulged with four lateral, oblique ridges. Pro-
notum transversely, finely rugulose. Scutel-
lum longer than pronotum.

Male genitalia. — Pygofer roundish with
macrosetae along caudal margin; ventral pro-
cess broad basally, abruptly narrowed near
apex to mesally directed, pointed process.
Anal tube moderately long. Male plate more
than four times as long as wide, single row of
macrosetae on caudal 0.75. Style with small,
preapical lobe, apophysis stout with short,
relatively stout apical extention. Connective
Y-shaped with stem longer than arm. Aedea-
gus flattened, elongate with lateral, marginal,
poorly sclerotized ridge, dorsal apodeme very
short, gonopore large, on caudal margin, sur-
rounded bv membranous extension caudallv.

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193

Figs. 49-68. Species ofKana: K. bispinosa, n. sp.: 49, head and thorax, profile; 50, same, dorsal view; 51, face; 52,
male pygofer; 53, style; 54, connective; 55, 56, aedeagus, lateral and ventral views; 57, male plate; 58, forewing; 59,
hindwing. K. nigropicta, n. sp.: 60, head and thorax; 61, forewing; 62, male pygofer; 63, apex of ventral process of
pygofer in two views; 64, style; 65, connective; 66, male plate; 67, 68, aedeagus, lateral and caudal views.

194

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Female genitalia. — Hind margin of sev-
enth sternum almost straight. Ovipositor ex-
ceeding pygofers.

Measurements. — Male 4.35 (4.20-4.50)
mm long, head 0.92 (0.90-0.95) mm wide,
pronotum 0.95 (0.90-0.95) mm wide. Female
5.13 (5.00-5.20) mm long, head 1.03
(1.00-1.08) mm wide, pronotum 1.11
(1.08-1. 15) mm wide.

Specimens examined. — Holotype 6, In-
dia: Kerala: Thekkadi, 26.iii.1977, C. A. Vi-
raktamath Coll. (UAS). Paratypes: 5 d, 2 9,
data as in holotype but collected on 27. iii. 1977
by C. A. Viraktamath (2 d, 1 9) (BMNH,
USNM, UAS).

Remarks. — This species can be easily rec-
ognized by the round, black spot on the vertex
and the absence of processes on the membra-
nous tube surrounding the gonopore.

Kana illaborata Distant

Figs. 69-79

Kana illaborata Distant 1908g: 287. Lectotype 3 , Burma
(BMNH, examined).

Yellow. Vertex with median, white stripe.
Pronotum with narrow, median, irregular,
lateral spot whitish. Forewing with broad,
median, mottled, brown band on apical 0.33.

Head about as wide as pronotum (46:47).
Vertex about as long as wide in male (29:28)
but longer than its width in female (38:34).
Pronotum shorter than both vertex and
scutellum. Face with four lateral, oblique
ridges and an apical, short, median ridge.

Male genitalia. — Pygofer with lobelike,
membranous process, armed with tooth
mesally; caudal margin with series of hairlike
setae. Male plate with oblique row of stout
setae in caudal half, and with marginal, hair-
like setae. Preapical lobe of style stout, apoph-
ysis relatively slender, apical extension slen-
der. Connective Y-shaped, with stem as long
as arm. Aedeagus laterally compressed, mem-
branous for the most part, with pair of slender
ventral processes and sclerotized tooth ven-
tral to gonopore, which is elongate; pair of
lamellate processes located in apical half of
shaft.

Measurements. — Male 5.70 mm long,
head 1.15 mm wide, pronotum 1.17 mm
wide. Female 6.30 mm long, vertex 1.37 mm
wide, pronotum 1.42 mm wide.

Specimens examined. — Lectotype 6 la-
beled /Myitta (Doherty)/ /Distant Coll.

1911-383/ /Kana illaborata Dist., det. M.
Webb, 1980/ here designated (BMNH).
Paralectotype 9 labeled /Tenass. Vail. Myitta
(Doherty)/ /Distant Coll. 1911-383/]Vtrt;ana
illaborata Dist. type/ here designated
(BMNH).

Remarks. — ^This species is very distinctive
both in coloration and male genitalic charac-
ters and does not appear closely related to
other species of Kana.

Kana decora (Melichar)

Figs. 80-81

Nirvana decora Melichar 1903b: 166. Holotype 9 , Sri
Lanka (ZMHU, examined).

The holotype female is a teneral specimen,
and the diagnostic coloration is not very well
developed. Melichar (1903b) has given a good
description of this species and hence it is not
redescribed here.

Specimen examined — Holotype 9, la-
beled /Type/ /6178/ /Ceylon, nictus/ /Nirvana/
/decora m. det. Melichar/ /Nirvana decora
Melichar/ (ZMHU).

Kana fasciata Pruthi

Kanafasciata Pruthi 1930a; 22. Syntypes d , 9 , Sri Lanka
(ZSI, not examined).

The types of this species were not available
for study at the time the senior author visited
ZSI. Pruthi's (1930a) description and illustra-
tions suggest this to be a species with very
distinctive coloration.

Genus Chudania Distant

Chudania Distant 1908g: 268. Type species Chudania
delecta Distant, by original designation and mono-
typy.

Yellow, with prominent broad stripe run-
ning on head, thorax, and along inner margin
of clavus, extending laterally to occupy entire
apex of forewing, black.

Vertex either as long as or slightly shorter
than width, disc convex without median
groove. Anterior area of disc rugose. Ocelli in
front of and mesad of eyes. Frontoclypeus
convex, with four to six transverse, prominent
rugae evanescent medially on dorsal aspect
and with median ridge in upper 0.75. Lora
prominent. Clypellus large, slightly narrowed
apically. Pronotum twice as wide as long,
slightly longer than scutellum, polished and
transversely, finely rugulose. Forewing with

1988

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195

tho'Sfe^l^trf '^pS.^r -^'^■' '': -r' ^™^''^^ ''- --' ^-- ^2' head and

79, .a,e forewing'. .. ^.^^ (tuchT^^^^^^^^^^ -^-^-> lateral and caudal views;

196

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No. 12

appendix wanting, apical cells four and an-
teapical cells two. Hindwing with four apical
cells. Hind tibial spinulation Rj 18 ±2, Ro
12±1, R3l2±l.

Male pygofer with rounded, caudal margin,
length shorter than height, strongly sclero-
tized with single, submarginal row of long
setae and ventral process. Anal tube stout and
large, about 0.8 times as high as height of
pygofer. Male plate with uniseriate macrose-
tae in apical 0.66 and with long, hairlike setae.
Style with prominent preapical lobe, apophy-
sis slender, apex avicephaliform. Style rather
T-shaped. Aedeagus membranous for most
part, bent in midlength, laterally expanded,
with ventrally directed, subapical, pronglike,
sclerotized processes surrounding gonopore.

Remarks. — This genus is related to Kana
and Afrokana Heller with which it shares
hindwing venation; with Afrokana it shares
the structure of the male pygofer. However,
they differ in the shape of the head. Lin-
navuori (1979) stated that A/roniruana Evans,
Chudania, and Afrokana are very close,
sharing, for instance, the same unique genital
structure. He believes the recorded differ-
ences are too slight for separating them as
valid genera. Afrokana is closer to Kana than
to Chudania considering the peculiar shape of
the aedeagus they share. It is likely that the
study of more material of these genera, espe-
cially from Malaysia, Indonesia, and the
Philippines, may prove that these taxa are
congeneric with Chudania.

Chudania delecta Distant

Figs. 82-96

Chudania delecta Distant 1908g: 268. Holotype 9 , India
(BMNH, examined).

Male. Slender, darker than female. Vertex,
pronotum, scutellum, upper part of face at
level with lower margins of eyes, chocolate
brown. Ocelli white, eyes dark brown.
Forewing yellow, with posteriorly widening,
black stripe along inner margin, occupying
entire apical 0.33 of forewing (Fig. 82); two
white, hyaline, triangular areas on costal area
near apex and some cross-veins enclosing api-
cal cells paler. Lower half of face, sterna, and
legs yellow. Pygofer and fifth to eighth terga
blackish.

Female. Similar to male but paler. Black
markings of male, chocolate brown. Entire

face, lateral margins of vertex in front of eyes,
and pronotum yellow.

Male genitalia. — Pygofer rounded cau-
dally, with submarginal row of macrosetae,
ventral process broad caudodorsally with
finely serrated, dorsal margin. Male plate
more or less of uniform width, caudally
rounded, with outer, marginal, long, hairlike
setae. Anal tube large, about as long as length
of pygofer. Style with anterior part larger than
posterior part to the point of articulation with
connective. Stem of connective thrice as long
as arms, apex expanded. Aedeagus membra-
nous for most part, processes sclerotized,
bent about its midlength, apical half robust,
preatrium with short ventral and longer lat-
eral processes; two pairs of processes arising
where shaft bends, dorsal pair elongate,
curved at midlength, the ventral pair shorter,
three to five branched.

Female genitalia. — Hind margin of sev-
enth sternum straight. Ovipositor exceeding
length of pygofer. Distal half of pygofer black.

Measurements. — Male 4.62 (4.60-4.70)
mm long, head 1.07 mm wide, pronotum 1.11
(1.07-1.12) mm wide. Female 5.38 (5.20-
5.40) mm long, head 1.20 mm wide, prono-
tum 1.22 (1.20-1.25) mm wide.

Specimens examined. — Holotype 9 la-
beled /Type, H.T./ /Chudania delecta Dist.,
type/ /Kurseong/ Distant Coll. 1911-383/
(BMNH). India: West Bengal: 8 d, 7 9,
Kurseong, 1,483 m, 22.x. 1981, C. A. & S.
Viraktamath; 1 9, 15km E Kahmpong, 1,780
m, 27.x. 1981, C. A. v.; 4 c?, 11 9, Kalimpong,
1,768 m, 29.x. 1981, S. V. ; 9 c? , 19 9 , 8 km E of
Kalimpong, 1,968 m, 29.x. 1981, C. A. V. Mi-
zoram: 11 d, 11 9, Aizawl, 18.xi.l981, C. S.
Wesley; 11 c?, 14 9 , Lungleh, 20-25. xi. 1981,
C.S.W. (UAS).

Remarks. — Chudania delecta and C. afri-
cana Heller are closely related but differ in
coloration, shape of the ventral pygofer pro-
cess, and the processes on the aedeagus. Also,
the dorsal half of the aedeagus is more robust
in C. delecta than in C. africana. The male
genitalia of the third species, C. exposita Ja-
cobi, is not known.

Genus Sophonia Walker

Sophonia Walker 1870b: .327. Type species: Sophonia

rufitehim Walker by monotypy.
Pseudonirvana Baker 1923a: 386. Type species:

Pseudonirvana sandakanensis Baker, by original

designation.

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ViRAKTAMATH, WESLEY; REVISION OF INDIAN NiRVANINAE

197

Figs 82-96 Chudania delecta Distant: 82, habitus, male; 83, head and thorax, male; 84, face, male; 85, head and
thorax, profile, male; 86, same, female; 87, forewing; 88, hindwing; 89, ovipositor; 90, male pygofer; 91, ventral process
of pygofer; 92, style; 93, 94, aedeagus, lateral and dorsal views; 95, connective; 96, male plate.

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No. 12

Quercinirvana Ahmed & Mahmood 1970: 260. Type spe-
cies: Quercinirvana longicephala Ahmed & Mah-
mood, by original designation. New synonymy.

Head as wide as or wider than pronotum.
Vertex as long as or longer than width be-
tween eyes, anterior half rugulose or granu-
lar, disc either flattened or slightly convex.
Ocelli lateral in front of eyes. Frontoclypeus
depressed, with prominent, lateral ridges and
with median, apical ridge; face longer than
width including eyes. Forewing with second
apical cell narrowed apically. Hindwing with
three apical cells. Hind tibial spinulation R,
20±2, R2l2±l, R3+ 1, R4l8±2.

Pygofer rounded or obtusely angled cau-
dally, with or without caudal or ventral pro-
cess; macrosetae confined to caudal 0.33 area.
Anal tube prominent. Male plate broader
than in Nirvana, three to four times as long as
its width and rather parallel-sided with
oblique row of stout spines in caudal 0.66 area
and hairlike setae. Connective Y-shaped, its
stem usually more than 1.5 times as long as
each arm. Aedeagus with dorsal apodeme well
developed, often with processes; preatrium
variable, with or without processes; shaft
tubular or lamellate, with or without process.

Remarks. — Evans (1947a) suppressed
Pseudonirvana as a junior synonym of S apho-
nia. This genus includes brightly colored spe-
cies, often with red or orange streaks. It is
closely related to Kana and Nirvana. It can be
diffierentiated mainly by its shorter male
plates and complex aedeagus that has pro-
cesses on the dorsal apodeme and apex of the
shaft. The holotype oi Sophonia rnfitelum is a
female (Figs. 97-102). The relationships dis-
cussed here are based on species assigned to
this genus from the Indian subcontinent.

Key to Species o( Sophonia

1. Vertex as long as wide between eyes (Figs. 186,

198) 2

— Vertex longer than wide between eyes (Figs.
112, 208) ' 3

2(1). Male pygofer with long, ventral and short, dor-
sal process (Fig. 203); preatrium without pro-
cesses bifida, n. sp.

— Male pygofer without processes; preatrium

with a pair of processes (Fig. 191)

modesta (Distant)

3(1). Vertex and pronotum with either single or
paired, black, longitudinal stripe(s) usually
connected to black spot or spots near apex of
vertex (Figs. 102, 1 12, 122) 4

— Vertex and pronotum without black, median
stripe or spot near apex; vertex sometimes with
median, bright crimson streak (Figs. 164, 177)

or orange stripe (Fig. 154) 8

4(3). Vertex with two rather oval, contiguous spots,
each connected to longitudinal, black line
(Figs. 103, 104); aedeagal shaft bifid (Figs. 109,
110) linealis (Distant)

— Vertex with a large, round, black spot at apex
either connected to or short distance away from
one or paired longitudinal line(s) (Figs. 112,
122); aedeagal shaft not bifid 5

5(4). Male pygofer without ventral or caudal process
(Fig. 125); aedeagus as in Fig. 129; median,
black lines on vertex not connected to its api-
cal, round spot (Fig. 122)

longitudinalis (Distant)

— Male pygofer with ventral or caudal process;
aedeagus variable but not as above (Figs. 118,
150); median, black stripe(s) of vertex con-
nected to spot at apex (Figs. 112, 132) 6

6(5). Vertex with single, median, black stripe (Fig.
142); male pygofer process short and narrow,
apophysis of style hooklike; apex of each arm of
dorsal apodeme with two long and short, me-
dian processes (Fig. 150) keralica, n. sp.

— Vertex with two median, black stripes (Fig.
112); male pygofer process broad and long; apo-
physis of style beaklike; each arm of dorsal
apodeme with a process on caudal margin (Fig.
118) 7

7(6). Male pygofer process with serrated, dorsal
margin; process of dorsal apodeme exceeding
width of shaft; shaft with pair of long, caudally

directed processes (Fig. 118)

longicephala (Ahmed & Mahmood)

— Male pygofer process not serrate apically; pro-
cess of dorsal apodeme not exceeding width of
shaft; processes of shaft short with basal, short,
ventral tubercle (Figs. 139, 141) . . bakeri, n. sp.

8(3). Male pygofer without process; preatrium of

aedeagus with process (Figs. 171, 180) 9

— Male pygofer with process (Figs. 159, 214);
preatrium of aedeagus without process 10

9(8). Preatrium longer than broad in lateral aspect;
longer process of dorsal apodeme not divided
(Fig. 171); vertex with short, median, crimson
streak in apical 0.66 but not reaching apex . . .
insignis (Distant)

— Preatrium broader than long, longer process of
dorsal apodeme bifid (Fig. 180); vertex uni-
formly whitish yellow with median, crimson
streak bifid near apex (Fig. 175) . complcxa, n. sp.

10(8). Vertex with median, broad, orange stripe con-
tinued on pronotinn, anteriorly narrowed; pos-
terior half on pronotum broadened and black-
i.sh, scutellum blackish (Figs. 154, 155);
pygofer process short, bladelike, and pointed
at apex (Fig. 159); aedeagus shaft tubular with
apical, ventrally directed, platelike process
(Figs. 162, 163) illuminata (Distant)

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ViRAKTAMATH, WESLEY: REVISION OF INDIAN NiRVANINAE

199

Figs. 97-102. Sophonia rufitehnn Walker: 97, head and thorax; 98, face; 99, head and thorax, profile; 100, hindwing;
101, basal half of clavus; 102, ovipositor.

— Vertex, pronotuni, and scutellum whitish yel-
low; pygofer process long, slender, and el-
bowed (Fig. 214); aedeagus not as above (Fig.
218) complicata, n. sp.

Sophonia linealis (Distant)
Figs. 103-111

Nirvana linealis Distant 1908g: 282. Holotype 9 , India

(BMNH, examined).
Nirvana greeni Distant 1908g: 283. Holotype 9 , Sri

Lanka (BMNH, examined). New synonymy.

Yellow. Vertex with two piceous, apical,
elongate, fused spots from which longitudi-
nal, piceous lines traverse posteriorly meet-
ing posterior margin of vertex, lateral margin
often with orange-yellow stripe confined to
ocelli or extending anteriorly. Pronotum and
scutellum with median, longitudinal, piceous
line often interrupted before apex of scutel-
lum. Claval margin of forewing piceous with
piceous stripe bent obliquely near claval apex

and reaching claval suture; spot on second
apical cell and two oblique lines from costa in
apical half piceous.

Head about as wide as pronotum. Vertex
more than 1.5 times as long as wide in male,
nearly twice as long as wide in female, lateral
margin raised. Scutellum longer than prono-
tum. Second apical cell of forewing narrowed
apically.

Male genitalia. — Pygofer ovate, without
ventral process. Plate slightly more than 3.5
times as long as median width, slightly nar-
rowed caudally. Style with broad, preapical
lobe and beaklike extension of apophysis rela-
tively short. Stem of connective flared cau-
dally, about twice as long as each arm. Dorsal
apodeme of aedeagus robust, each arm with
caudally directed, elongate process that ex-
ceeds width of shaft; shaft tubular, curved,
hooked apically where it appears bifid.

200

Great Basin Naturalist Memoirs

No. 12

Figs. 103-121. Sophonialinealis {Distant): 103, head and thora.x of holotype female Niruaria greeni Dist.; 104, head
and thorax of holotvpc of Nirvana linealis Dist.; 105, profile; 106, ovipositor; 107, male pygofer; 108, forewing; 109
110, aedeagus, lateral and dorsal views; HI, connective and style. S. km^icephala {Ahmed in Mdhmood): 112, head and
thorax; 113, face; 114, male plate; 115. male pygofer; 116, ventral process of pygofer; 117, style; 118, 119, aedeagus,
lateral and caudal views; 120, connective; 121, forewing.

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ViRAKTAMATH, WeSLEY: REVISION OF INDIAN NiRVANINAE

201

Female genitalia. — Hind margin of sev-
enth sternum straight with median protuber-
ance.

Measurements. — Male 4.06 (4.00-4.10)
mm long, head 0.83 (0.80-0.85) mm wide,
pronotum 0.88 (0.85-0.93) mm wide. Female
4.82 (4.50-5.00) mm long, head 0.93
(0.88-0.95) mm wide, pronotum 0.98
(0.93-1.03) mm wide.

Specimens examined. — Holotype 9, la-
beled/Type, H.T./ZPeradeniya, 2.08, Ceylon/
/Distant Coll. 1911-383/ /Nirvana greeni
Dist., type/ (BMNH). Holotype 9, labeled
/Type, H.T./ /Calcutta, 6.6.07/ /Distant Coll.
1911-383/ /Nirvana linealis Dist., type/
(BMNH). India: Karnataka: 6 9, Dharwar,
26.iii.1972; 1 9 29. iv. 1969; 1 9, — .i.l970; 1
9, 14. iv. 1972; 1 9, — .x.l979 C. A. Virakta-
math Coll.; 3 d, 3 9, Chamarajanagar,
12.viii.l977; 4 9, Hunsur, 16.i.l978; 1 9,
Bangalore, 11. vi. 1977; 1 9, Arsikere,
23.vii.1978; 1 d, 1 9, 16.i.l979; 1 9, Sulik-
ere, 30.xii.l976, all collected by C. A. Virak-
tamath (UAS). Kerala: 3 c? , 6 9 , Kayangulum,
— .viii. 1983, ex coconut (UAS).

Remarks. — The holotypes of linealis and
greeni show color differences, especially the
extension of the median, dark fuscous stripe of
the pronotum along the inner margin of the
clavus in greeni, which is absent in linealis.
The female seventh sterna are identical;
hence the species are considered as syn-
onyms. This species is unique among Sopho-
nia in having a bifid aedeagal shaft.

Sophonia longicephala

(Ahmed & Mahmood), n. comb.

Figs. 112-121

Quercinirvana longicephala Ahmed & Mahmood 1970:
263. Holotype <5, Pakistan (University of Karachi,
not examined).

Pale yellow, elongate, black spot near apex
of vertex contiguous with two longitudinal,
black lines that fuse at base of vertex and
continue as single line to near tip of scutellum.
Forewing with broad, brown stripe along
claval commissure reaching claval apex and
obliquely crossing corium, apical margin suf-
fused with brown, costal margin with ante-
rior, oblique and two posterior, transverse,
black fasciae; second apical cell with black
spot.

Head nearly as wide as pronotum. Vertex
about 0.33 times longer than wide, disc de-

pressed, faintly granulose. Second apical cell
narrowed caudally.

Male c;enitalia. — Pygofer trapezoidal,
with long, caudally direct(^d ventral process
dorsally serrated. Plate about four times as
long as wide, distal 0.66 with a single row of
macrosetae and hairlike setae. Style with
large, preapical lobe, beaklike extension of
apophysis long. Aedeagus with dorsal
apodeine well-developed, caudally directed,
apically pointed processes that exceed width
of shaft, shaft tubular, dorsoanteriorly curved
with subapical, caudally directed, slender,
elongate processes.

Measurements. — Male 4.50 (4.30-4.70)
mm long, head 0.91 (0.90-0.93) mm wide,
pronotum 0.96 (0.93-1.00) mm wide.

Specimens examined. — India: Haryana: 1
6, Kalka, 29.i.l979, C. A. Viraktamath; Hi-
machal Pradesh: 1 6, Kulu, l.iv.l978, I.
Dworakowska (UAS).

Remarks. — This species is related to and
resembles externally S. bakeri. The spot on
the vertex in this species is rather oval (as in
bakeri), and it differs in the details of the male
pygofer process and the aedeagus.

Sophonia longitudinalis (Distant)
Figs. 122-1.31

Nirvana longitudinalis Distant 1908g: 283. Lectotype 9 ,
Burma (BMNH, examined).

Vertex pale yellow with large, subapical,
black spot and two median, longitudinal lines,
not reaching black spot and fused near hind
margin of vertex, continued on pronotum and
scutellum as median stripe and on clavus as
irregular stripe. Forewing with large, round,
black spot in second apical cell extending
partly to first apical cell, apical margin suf-
fused with brown.

Head as wide as pronotum or slightly nar-
rower. Vertex about 1.5 times as long as wide,
apex subangular, disc mildly convex, granu-
lose. Second apical cell narrowed caudally.

Male genitalia. — Pygofer ovate, ventral
process wanting, macrosetae limited to distal
0.33. Plate about 3.5 times as long as width,
with short tooth at the outer apical margin.
Style with short, preapical lobe, apophysis
with long, beaklike extension. Aedeagus with
robust, dorsal apodeme, each of its arms bear-
ing an elongate, caudally directed process,
preatrium wanting, shaft lamellate with lat-
eral margin caudally curved, widest at apical

202

Great Basin Naturalist Memoirs

No. 12

Figs. 122-141. Sophonialongitudinalis (Vistsint): 122, head and thorax; 123, face; 124, ovipositor; 125, male pygofer;
126, 127, male plate; 128, connective and stvle; 129, 130, aedeagus, lateral and caudal views; 131, forewing. S. I^^ken,
n. sp.: 132, head and thorax; 133, face; 134, forewing; 135, male pygofer; 136, pygofer process; 137, male plate; 138,
connective; 139-141, aedeagus, lateral, caudal, and dorsal views.

1988

ViRAKTAMATH, WeSLEY: REVISION OF INDIAN NiRVANINAE

203

half with long, slender, arcuate, apically
pointed, ventrally curved process that bears a
short tooth on its outer margin at base.

Female genitalia. — Hind margin of sev-
enth sternum slightly concave. Ovipositor
slightly exceeding pygofer.

Measurements— Male 4.43 (3.80-5.40)
mm long, head 0.96 (0.80-1.43) mm wide,
pronotum 0.96 (0.85-1. 13) mm wide. Female
4.90 (4.60-5.30) mm long, head 1.02
(0.98-1.10) mm wide, pronotum 1.05
(1.00-1. 10) mm wide.

Specimens examined. — Lectotype 9 la-
beled /Myitta (Doherty)/ /Distant Coll.
1911-383/ /Nirvana longitudinalis Dist.,
type/ here designated (BMNH). India: Karna-
taka: 6 (5, Mudigere, 4-6. iv. 1980, C. A. Vi-
raktamath; 1 9, 7.iv.l975, C.A.V.; 1 9,
Kodlipet, 6.iv.l975; 1 S, Hunsur, 10. i. 1978;
1 9, Bannerghatta, 12. ix. 1976, B. Mallik;
Kerala: 1 9, Thekkadi, 27.iii.1977, C.A.V.
Nepal: Nagarkot, 1 (5, 1 9, 15.ix.l979,- I.
Dworakowska (UAS). Burma: 1 9, Amberst
Dist., Kawkareik, 19-20.xi.l911, F.H.G.
(ZSI).

Remarks. — This species can be distin-
guished from all other species of Sophonia by
its lamellate aedeagal shaft and tooth on the
apex of the male plate.

Sophonia bakeri, n. sp.
Figs. 132-141

Pale yellow to whitish yellow. Vertex with
black spot near apex, joined by two black,
longitudinal lines that fuse basally and extend
to scutellum. Pronotum and scutellum some-
times with reddish spots, scutellum in male
entirely reddish. Forewing with brown stripe
close to or along anal margin, marked with
yellow along inner margin, apex of clavus and
apical margin brown, costal margin with ante-
rior, oblique and posterior, transverse, brown
fasciae, second apical cell with large, black
spot.

Vertex about 1.5 times as long as wide, apex
subtriangular, disc depressed. Posterior half
of pronotum transversely rugulose. Second
apical cell narrowed at apex.

Male genitalia. — Pygofer with ventrocau-
dal angle produced to process with attenuated
apex. Plate 3.5 times as long as wide. Apophy-
sis of style with long, beaklike extension.
Stem of connective 1.5 times as long as each
arm. Aedeagus with each arm of dorsal apo-

deme robust, bearing triangular process on
caudal margin; preatrium short; shaft tubular,
caudodorsally curved, apex strongly incurved
with ventrally directed, short, pronglike pro-
cess on caudal margin subapically.

Female genitalia. — Hind margin of sev-
enth sternum concavely emarginate. Oviposi-
tor exceeding length of pygofer.

Measurements,— Male 4.48 (4.30-5.00)
mm long, head 0.90 (0.85-1.00) mm wide,
pronotum 0.88 (0.78-1.03) mm wide. Female
5.08 (4.70-5.30) mm long, head 1.04 (0.93-
1.23) mm wide, pronotum 1.05 (1.00-1.10)
mm wide.

Specimens examined. — Holotype 6, In-
dia: Karnataka: Mudigere, at light,
4-6.iv.l980, C. A. Viraktamath (UAS).
Paratypes: India: Karnataka: 2 6, Dharwar,
— .i.l970, C. A. Viraktamath; 1 9, Kodlipet,
6.iv.l975, C.A.V. ; 1 9, Bangalore, 9.iii.l975,
C.A.V.; 1 9, Kalhalla, 18.i.l978, C.A.V., 1
9, Mudigere, 23. ix. 1973, C.A.V.; 1 (?, 1 9,
Kemmanagundi, 8.iv.l975, K. D. Ghorpade;
Kerala: 1 c?, Thekkadi, 27.iii.1977, S. Virakta-
math; Maharashtra: 1 9 , Matheran, 915 m,
23.xi.1977, C.A.V.; Tamil Nadu: 1 6, Ko-
daikanal, 2,250 m, 10. vi. 1980, C. S. Wesley;
Uttar Pradesh: 1 6, 1 9, Dehra Dun,
28. iv. 1975, C.A.V.; Nepal: 2 9, Gokarna,
14. ix. 1979, I. Dworakowska; 1 9, Sarankot,
17. ix. 1979, exQiiercus sp., I.D. (1 6 and 1 9
in BMNH, lARI, USNM, and rest in UAS).

Remarks. — Sophonia bakeri is related to S.
longitudinalis and S. longicephala as they
share similar external coloration and male
genitalia. They form a closely knit group. S.
bakeri differs from the other two by its shorter
process of the dorsal apodeme and aedeagal
process.

Sophonia keralica, n. sp.
Figs. 142-153

Uniformly yellow. Vertex with round, black
spot near apex connected by single, median,
longitudinal, black line, discontinuous for
short distance near black spot and then con-
tinued to apex of scutellum. Lateral margin of
vertex orange basally. Forewing with claval
suture and punctures along cubital vein
brown, second and part of third apical cells
occupied by large, black spot; costal margin
with anterior, oblique and posterior, trans-
verse, black fasciae, apex of clavus and apical
margin brownish.

204

Great Basin Naturalist Memoirs

No. 12

Figs. 142-153. Sophonia keralica, n. sp.: 142, head and thorax; 143, face; 144, head and thorax, profile; 145, male
pygofer; 146, pygofer process; 147, connective; 148, style; 149, male plate; 150, 151, aedeagus, lateral and cephalic
views; 152, forewing; 153, hindwing.

Head nearly as wide as pronotum. Vertex
slightly longer than wide, disc flat, depressed
near apex. Scutellum slightly longer than
pronotum. Second apical cell of forewing nar-
rowed slightly near apex.

Male genitalia. — Pygofer caudally
rounded, its ventrocaudal angle produced to
sharp, thin process, five macrosetae along
caudal margin in straight line, black, rest hya-
line. Anal tube as long as pygofer. Plate four
times as long as wide. Style with strongly
hooked apophysis. Stem of connective 1.5
times as long as each arm. Dorsal apodeme of
aedeagus slender, each arm with ventral, dor-
sal, and short, median process; shaft tubular
with pair of laterally directed processes,
gonopore subapical.

Measurements. — Male 3.90 mm long,
head 0.90 mm wide, pronotum 0.93 mm
wide.

Specimen examined. — Holotvpe 6, India:
Kerala: Thekkadi, 27.iii.1977, s! Viraktamath
Coll. (UAS).

Remarks. — Sophonio keralica is related to
S. iUuminata. Both have similar male geni-
talia. They differ, however, in coloration and
detailed structure of the aedeagus.

Sophonia iUuminata (Distant), n. comb.
Figs. 154-163

Kana iUuminata Distant 1918b: 35. Lectotype d, India

(BMNH, examined).
Kana sigruiffl Distant 1918b: 34. Lectotype 9 , India

(BMNH, examined). New synonymy.

Female. Vertex and pronotum creamy
white with yellowish lateral areas, median
stripe narrowing to apex of vertex san-
guineous on vertex and anterior half of prono-
tum, widened in posterior half where it is
black. Scutelhun black except for two lateral,
basal spots. Clavus along claval suture and
apical half piceous except for subapical, elon-
gate, orange spot. Corium along its inner mar-
gin piceous which widens near apex of cla-
vus and changes to orange; appendix fuscous,
two large, round spots in second apical cell.

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205

Figs. 154-163. Sophonia illuminata (Distant): 154, head and thorax of lectotype male Kana illuminata; 155, head and
thorax of lectotype female Kana signata Distant; 156, forewing, female; 157, ovipositor; 158, apex of male pygofer; 159,
pygofer process; 160, style; 161, connective; 162, 163, aedeagus, lateral and caudal views.

oblique line at apical 0.33 of costa and more
distal, transverse band piceous, rest of wing
yellow and areas anterior to piceous spots
whitish.

Male. Coloration less extensive compared
to female. Stripe on vertex continuing on
pronotum and scutellum, gradually widening
and sanguineous throughout. Forewing col-
oration comparatively less developed.

Head narrower than pronotum, median
length of vertex slightly longer than width
between eyes. Face longer than wide.

Male genitalia. — Pygofer lobe gradually
narrowed caudally and terminated by sharp,
triangular process and with series of stout,

long spines. Plate parallel-sided, with six
stout setae in oblique row commencing at
midlength, third to fifth setae black in color.
Style with stout, preapical lobe and hooklike,
rather slender apophysis with attenuated
apex. Connective with stem 1.5 times as long
as each arm. Aedeagus with short, dorsal pair
of blunt processes and ventral, fingerlike pro-
cess; shaft robust at base, narrowed apically
with dorsal, large gonopore surrounded by
two lateral, pronglike processes and ventrally
directed, lobelike plate.

Female genitalia. — Hind margin of sev-
enth sternum concave. Ovipositor exceeding
length of pygofer.

206

Great Basin Naturalist Memoirs

No. 12

Measurements. — Male 4.40 mm long,
head 1.00 mm wide, pronotum 1.05 mm
wide. Female 5.1 mm long, head 1.12 mm
wide, pronotum 1.12 mm wide.

Specimens examined. — 3 syntype 6 of
Kana illuminata mounted on single card la-
beled /S. India, Madras, Kodaikanal, T. V.
Campbell, K.K.4.16/. One of these was re-
moved, remounted, and relabeled by the se-
nior author, /Kodaikanal, S. India, T. V.
Campbell/ and is here designated as lecto-
type; others designated paralectotypes. One
paralectotype 6 labeled /Kodaikanal, India,
T. V. Campbell/ /S. India, E. A. Butler,
1915-60/ /Kana illuminata Dist., type/
(BMNH). Lectotype 9 labeled /Kodaikanal,
S. India, T. V. Campbell, K.K.5.11//S. India,
E. A. Butler, 1915-60/ /54,9/ /Kana signata
Dist., type/ here designated (BMNH). India:
Tamil Nadu: 1 9, Trichinopoly, J. Dubre-
nil(IRSNB); 1 9 Kadaikanal, 2,030-2,150 m,
— .viii.1922, S. Kemp(ZSI).

Remarks. — Sophonia illuminata and S. sig-
nata are the male and female, respectively, of
the same species based on the similarity in
external coloration and structure. This species
is closely related to S. keralica. Both have a
similar type of male genitalia but differ in
coloration and structure of the pygofer pro-
cess and aedeagus.

Sophonia insignis (Distant), n. comb.
Figs. 164-174

Nirvana insignis Distant 1918b: 33. Lectotype 6, India
(BMNH, examined).

Whitish yellow. Median groove on vertex
crimson from about center to just short of
apex, lateral marginal area orange. Forewing
hyaline, light brown stripe along anal margin
rarely reaching apex of clavus, if so occupying
entire quarter of clavus, or stripe wanting,
apex of clavus brown; costal margin with two
long, oblique fasciae, third transverse and
fourth short, broad, oblique fasciae in the dis-
tal half brown; apical margin brown; second
apical cell with black spot.

Head narrower than pronotum. Vertex
about 1.5 times as long as wide between eyes;
disc flat or slightly depressed toward base,
apex granulose. Second apical cell of forewing
slightly narrowed distally.

Male genitalia, — Caudal lobe of pygofer
rounded, without processes. Plate about 3.5
times as long as wide. Preapical lobe of style

robust, apophysis slender, with apical exten-
sion forming short, beaklike extension.
Aedeagus with preatrium well developed,
with ventrocaudally directed process, about
as long as preatrium; dorsal apodeme broad,
platelike, with two processes (one short,
pronglike on cephalic margin, other long,
slender, and inwardly curved in middle) and
ventral, short process; shaft tubular, curved
anteriorly then ventrally near apex.

Female genitalia. — Hind margin of sev-
enth sternum straight. Ovipositor slightly ex-
tending beyond pygofer.

Measurements— Male 5.00 (4.80-5.20)
mm long, head 0.97 (0.93-1.00) mm wide,
pronotum 1.04 (1.03-1.07) mm wide. Female
5.62 (5.00-6.10) mm long, head 1.06 (0.98-
1.13) mm wide, pronotum 1.14 (1.03-1.23)
mm wide.

Specimens examined. — Lectotype S la-
beled /Tenmalai, W. Ghat (W. side). Travan-
core, 22.xi.08, Annandale/ /Distant Coll.
1911-383/ here designated (BMNH). Paralec-
totype 9 , with same data as in holotype, here
designated (ZSI). India: Kerala: 11 (5, 6 9,
Thekkadi, 26-27. iii. 1977, C. A. Viraktamath,
S. Viraktamath, and B. Mallik Coll.; Kar-
nataka: 2 9, Mudigere, 7.iv.l980, C.A.V.
(UAS).

Remarks. — This species is closely related
to S. complexa and can be differentiated by
the structure of male genitalia and coloration.

Sophonia complexa, n. sp.

Figs. 175-185

Similar to S. insignis but more elongate.
Vertex with median, crimson-colored line oc-
cupying about 0.66 area and apically bifid,
short, slightly oblique, sublateral stripe on
apical half of vertex orange. Forewing with
fuscous spots (Fig. 178).

Apical 0.75 of vertex elongately rugulose,
disc flat, margined by carina.

Male genitalia. — Pygofer elongate, heav-
ily setose, caudally rounded. Anal tube stout,
long. Plate parallel-sided, about 3.5 times as
long as wide, macrosetae restricted to apical
0.66. Apophysis of style short, apex avicephal-
iform. Connective with stem about three
times as long as each arm. Aedeagus complex,
each arm of dorsal apodeme with pair of ven-
trally directed processes, preatrium with cau-
dally directed process, shaft tubular, slightly
curved dorsally near apical 0.33, slightly
flared.

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207

173^ 170 174

Figs. 164-174. Sophonia insignis (Distant): 164, head and thorax; 165, face; 166, ovipositor; 167, male pygofer; 168,
style; 169, connective; 170, male plate; 171-173, aedeagus, lateral, cephalic, and caudal views; 174, forewing.

Female genitalia. — Hind margin of sev-
enth sternum slightly concave.

Measurements.— Male 5.32 (5.20-5.50)
mm long, head 1.00 (0.97-1.02) mm wide,
pronotum 1.06 (1.05-1.07) mm wide. Female
5.92 (5.80-6.00) mm long, head 1.10 (1.05-
1.12) mm wide, pronotum 1.16 (1.15-1.17)
mm wide.

Specimens examined. — Holotype 6, In-
dia: Meghalaya: Nongpoh, 762 m, 4.xi.l981,
C. A. Viraktamath (UAS). Paratypes: 13 d, 4
9 , data as in holotype but 7 d , 2 9 S. Virakta-
math Coll. (1 d and 1 9 in BMNH, lARI,
USNM, and rest in UAS).

Remarks. — This species is related to and
resembles S. insignis as they share coloration
and similar aedeagi. However, S. cornplexa
differs from S . insignis in having the process
of the dorsal apodeme bifid ventrally, a longer

stem on the connective, and the crimson line
of the vertex being forked apically.

Sophonia rnodesta (Distant), n. comb.

Figs. 186-197

Kana rnodesta Distant 1918b: 36. Lectotype 6, India
(BMNH, examined).

Pale yellow. Vertex of female creamy
white, eyes lemon yellow. Forewing hyaline
with brown stripe close to anal margin often
reaching claval apex, apex of corium lemon
yellow; costal margin with an anterior, long,
oblique, median and a slightly oblique, poste-
rior, transverse, brown fasciae; second apical
cell with black spot, apical margin brownish.

Head slightly broader than pronotum. Ver-
tex as long as wide between eyes, disc convex,
apex subangular, granulose.

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Figs. 175-185. Sophonia complexa, n. sp.: 175, head and thorax; 176, same, profile; 177, face; 178, forewing; 179,
pygofer; 180, 181, aedeagus, lateral and caudal views; 182, style; 183, male plate; 184, connective; 185, ovipositor.

Male genitalia. — Caudal lobe of pygofer
rounded, without process. Anal collar well
developed. Plate four times as long as wide.
Preapical lobe of style small, rounded, apoph-
ysis setose on lateral margin, apical extension

beaklike. Stem of connective four times as
long as each arm. Aedeagus with preatrium
short, pair of ventrocaudally directed pro-
cesses, dorsal apodeme robust, each arm
rounded; shaft tubular, slender, dorsally

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ViRAKTAMATH, WESLEY. REVISION OF INDIAN NiRVANINAE

209

Figs. 186-197. Sophonia modesta (Distant); 186, head and thorax; 187, same, profile; 188, face; 189, ovipositor; 190,
pygofer; 191, aedeagus and part of connective, lateral view; 192, 193, aedeagus, dorsal and cephalic views; 194, style;
195, connective; 196, male plate; 197, forewing.

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strongly curved anteriorly, apex ventrally di-
rected with pair of elongate, basal processes
that are directed dorsoanteriorly then later-
ally.

Female genitalia, — Hind margin of sev-
enth sternum slightly convex. Ovipositor fair-
ly exceeding pygofer.

Measurements. — Male 4.43 (4.40-4.50)
mm long, head 1.08 (1.07-1.10) mm wide,
pronotum 1.05 (1.03-1.08) mm wide. Female
5.20 (5.00-5.40) mm long, 1.24 (1.20-1.28)
mm wide, pronotum 1.19 (1.15-1.25) mm
wide.

Specimens examined. — Lectotype S la-
beled /South India, Madras, Coonoor, T. V.
Campbell/ here designated (BMNH). India:
Tamil Nadu: 3 (5 , 1 9 , Kotagiri, 23.x. 1975, C.
A. Viraktamath; 1 9, Coonoor, ll.viii.l979,
S. Viraktamath; 1 6, Wellington, 12.viii.
1979, S.V.; 5 9, Ootacamund, 2,350 m, 13.
viii.1979, S.V. (UAS).

Remarks. — Sophonia modesta externally
resembles S. bifida in having a wider vertex,
but they are only distantly related. It is re-
lated to S. insignis and S. complexa and differs
from them in having paired preatrial pro-
cesses and lacking processes on the dorsal
apodeme. In these three species the stem of
the connective is more than three times as
long as each arm.

Sophonia bifida, n. sp.

Figs. 198-207

Bright lemon yellow. Two spots on inner
claval margin, one at midlength, another at
apex, small costal patch about midlength, an
outer, oblique and posterior, transverse fascia
on costal area fuscous, large, round spot on
second apical cell black. Hind tibial apical
pecten black.

Head slightly narrower than pronotum.
Disc of vertex about as long as wide between
eyes, broadly rounded with apical area finely
rugulose, posterior half polished. Fronto-
clypeus rather tumid. Second apical cell of
forewing slightly narrowed caudally.

Male genitalia. — Caudal margin of
pygofer rounded with caudodorsal, short,
stout, strongly hooked process and caudoven-
tral, elongate, sharply pointed process;
macrosetae forming single row. Anal tube
large. Preapical lobe of style short, rounded,
apophysis slender, rather hooked, apex beak-
like. Arms of connective about as long as

stem. Aedeagus with elongate preatrium;
each arm of dorsal apodeme rounded; shaft
tubular with basal pair of elongate processes
extending to about 0.75 of length and with
subapical pair of denticles on caudal margin.

Measurements. — Male 4.50 mm long,
head 1.07 mm wide, pronotum 1.10 mm
wide. Female 5.30 mm long, head 1.25 mm
wide, pronotum 1.25 mm wide.

Specimens examined. — Holotype 6, In-
dia: West Bengal: Darjeeling, 2,176 m,
11. xi. 1981, C. S. Wesley (UAS). Paratype 9,
India: West Bengal: Kalimpong, 1,370 m,
29.x. 1981, S. Viraktamath (UAS).

Remarks. — Although the shape of the head
is similar to that in S. modesta, S. bifida occu-
pies an isolated place in Sophonia. The pecu-
liar pygofer process and aedeagus differenti-
ate it from all other species.

Sophonia complicata, n. sp.

Figs. 208-220

Whitish yellow. Area surrounding ocelli,
lateral-most area of pronotum lemon yellow.
Second apical cell with round, black spot.
Costa with outer, oblique fascia, posterior,
transverse fascia near apex, and apical margin
of forewing fuscous. In a few specimens, me-
dian area of clavus pale fuscous, which meets
fuscous stripe on claval suture and continues
angularly across wing where it is joined by an
oblique, fuscous fascia from costa. Hind tibial
pecten fuscous.

Vertex more conically produced, disc con-
vex with median carina in apical half and api-
cal area rugulose, rugae crisscrossing. Vertex
of female distinctly longer than male. Head
wider than pronotum.

Male genitalia. — Pygofer with caudodor-
sal angle roundly produced, with excavation
below which long, slender process arises, di-
rected caudally, upcurved at right angles at
midlength, apex slightly hooked, bearing
spinelike branch at base. Anal tube robust,
slightly exceeding pygofer lobe, but not
pygofer process. Plate 3.75 times as long as
wide. Style with elongate, slender, preapical
lobe, apophysis slender avicephaliform. Stem
of connective more than twice as long as each
arm. Aedeagus complex, two arms of dorsal
apodemes well separated, winglike, com-
pletely covering entire length of shaft later-
ally, each arm with dorsomedian, caudal, and
caudoventral, long processes; dorsomedian

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211

204

206

207

Figs. 198-207. Sophonia bifida, n. sp.: 198, head and thorax; 199, same, profile; 200, face; 201, forewing; 202, male
pygofer; 203, pygofer process; 204, style; 205, connective; 206, 207, aedeagus, lateral and caudal views.

process curves caudally, caudal process di-
rected anteriorly and in contact with dorso-
median process, caudoventral process crosses
over caudad of shaft, thus enclosing it; shaft
depressed, strongly curved anteriorly, ex-
panded, and hoodlike.

Measurements. — Male 4.40 (4.30-4.50)
mm long, head 0.92 (0.90-0.95) mm wide,
pronotum 0.94 (0.90-0.95) mm wide. Female
4.88 (4.60-5.10) mm long, head 1.01 (0.97-
1.05) mm wide, pronotum 1.02 (1.00-1.05)
mm wide.

Specimens examined. — Holotype 6, In-
dia: Mizoram: Lungleh, 23. xi. 1981, C. S.
Wesley (UAS). Paratypes: 15 (5, 16 9, India:
Mizoram: Aizawl, 18. xi. 1981, C. S. Wesley; 3
(5, 1 9, Lungleh, 22-24.xi.l981, C. S.
Wesley; Meghalaya: 2 9, Cherrapunji, 1,299
m, 3.xi.l981, C. A. Viraktamath; 1 9,
Shillong, 1,961 m, 3..xi.l981, S. Viraktamath

(1 d and 1 9 paratypes in BMNH, lARI,
USNM, ZSI, and rest in UAS).

Remarks. — The complex nature of the
aedeagus and the slender, long pygofer pro-
cess distinguish this species from other spe-
cies oi Sophonia.

Genus Nirvana Kirkaldy

Nirvana Kirkaldy 1900d: 293. Type species: Nirvana
pseudommatos Kirkaldy, by original designation
and monotypy.

Vertex longer than width between eyes,
disc depressed apically, with median ridge.
Ocelli situated laterally mesad of submarginal
carina. Face slightly depressed basally, fron-
toclypeus with short, apical, median ridge and
lateral, oblique rugae weakly developed. Sec-
ond apical cell of forewing narrowed apically.
Hindwing with three apical, closed cells.

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Figs. 208-220. Sophonia complicatd. n. sp.: 208, head and thorax; 209, same, profile; 210, face; 211, forewing; 212,
ovipositor; 213, male pygofer; 214, pygofer process; 215, style; 216, male plate; 217, connective; 218-220, aedeagus,
lateral, caudal, and dorsal views.

Hind tibial spinulation R, 20 ± 2. R. 12 ± 1, R,
12 ±2.

Male genitalia. — Pygofer lobe caudally,
bluntly conical or rounded, with ventral,
.short or long proccs.s, caudal area with numer-
ous macrosetae. Plate parallel-sided, elon-
gate, more than five times as long as wide near
middle, with oblique row of macrosetae in
apical 0.66 and hairlike setae. Style with well-
developed preapical lobe; apophysis short,
apical extension more than twice length,
beaklike. Aedeagal shaft tubular, with or
without processes, dorsal apodcme slender in
cephalic view, U-shaped; gonoporc apical.

Remarks. — Nirvana and So})honia are
closely related and are ditficult to separate

by external characters. Nirvana has a simple
dorsal apodeme, usually without processes,
whereas in Soplionia it is complex with more
elaborate processes; the male plate in Nirvana
is narrow and elongate compared to Sopho-
nia. Specimens oiN. psetido)n))iato.s were not
available for the study, and hence the genus is
only tentatively defined here.

Key to Species oiNirvana

I. Vertex with median, white stripe and lateral,
orange or lemon yellow, suhmarginal stripe;
aedeagal shaft with two slender, lateral, long,
ventral processes directed ventrally (Figs. 225,
226) pallida Melichar

— Vertex with black line or lines or with orange,
suhmarginal stripes traversing on pronotum and

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213

scutellum and forewing; aedeagus without pro-
cesses (male of pseudommatos is not known) ... 2

2(1). Vertex, pronotum, scutellum traversed by pair
of orange stripes (Fig. 256); male pygofer cau-
dally produced into an acute, dorsally curved
process (Fig. 261); anal tube with two stout
spines (Fig. 262) pecuhahs. n. sp.

— Vertex, pronotum, and scutelhun traversed by
black lines or stripes; male pygofer and anal tube
unarmed (not known in pseudommatos ) 3

3(2). Vertex, pronotum, scutellum, and anal margin

of forewing traversed by broad, black stripe .... 4

— Vertex of head and pronotum traversed by three
black stripes; scutellum by a median, black line;
apex of vertex with blackish brown spot con-
nected by the stripes .... pseudommatos Kirkaldy

4(3). Median stripe with irregular margins (Figs. 233,
234); spot on second apical cell of forewing small
(Fig. 243); dorsal apodeme of aedeagus simple
(Fig. 240) suturalis Melichar

— Median stripe with an even margin (Fig. 244);
spot on second apical cell of forewing large (Fig.
248) extending to first apical cell; dorsal
apodeme of aedeagus with a basal, short process
(Fig. 254) striata, n. sp.

Nirvana pallida Melichar

Figs. 221-232

Nirvana pallida Melichar 1903b: 166. Syntype cJ, 9, Sri
Lanka (not examined).

Quercinirvana bengalensis Ahmed & Mahmood 1970:
263. Holotype d, Bangladesh (University of
Karachi, not examined). New synonymy.

Coloration as described by Melichar
(1903b). The median, white stripe of vertex
may be obscured in some specimens.

Head as wide as or slightly narrower than
pronotum. Vertex 1.5 to 2.0 times as long as
width between eyes, disc slightly depressed.
Clypellus convex.

Male genitalia. — Pygofer lobe caudally
rounded with marginal macrosetae, ventral
process long, dorsally directed with rounded
apex. Plate narrowed caudally with bluntly
conical apex. Preapical lobe of style rounded,
stout. Stem of connective longer than arms.
Aedeagus with dorsal apodeme well devel-
oped, consisting of two arms and forming a U,
shaft tubular, curved slightly with short,
spinelike, laterally directed, terminal pro-
cesses and long, unpaired, ventrally directed,
subapical process.

Female genitalia. — Hind margin of sev-
enth sternum rather straight. Ovipositor
slightly exceeding pygofer.

Measurements.— Male 4.80 (4.60-5.00)
mm long, head 0.94 (0.90-0.98) mm wide.

pronotum 1.02 (0.95-1.08) mm wide. Female
5.65 (5.5-5.80) mm long, h(>ad 1.06 (1.05-
1.07) mm wide, pronotum 0.93 (0.88-0.98)
mm wide.

Specimens examined. — India: 203 d, 274
9 , collected from the states of Andhra
Pradesh, Haryana, Karnataka, Kerala, Maha-
rashtra, Meghalaya, Mizoram, Sikkim, Tamil
Nadu, and West Bengal from January to De-
cember, from sea level (Cochin in Kerala) to
2,250 m (Kodaikanal in Tamil Nadu) above sea
level. Nepal: 1 9, Sarankot, 17.xi.l979, I.
Dworakowska. Sri Lanka: 1 9 , Peradeniya,
23.viii.1979, I. Dworakowska (UAS).

Remarks. — This species is the most com-
mon and most frequently encountered Nir-
vaninae on the Indian subcontinent. It breeds
on newly emerged leaves of Tecoma stans,
Duranta sp., Vitis vinifera, beans, pigeon
pea, etc. The species is identified based on the
description and illustrations provided by
Melichar (1903b). No other species of Nirvan-
inae has the color and elongate male plate
with two short, lateral, and caudal processes
on the aedeagal shaft. The synonymy of Q.
bengalensis is based on the illustrations of
male genitalia of the species provided by
Ahmed and Mahmood (1970) and on the spec-
imens collected from northeastern India.
None of the specimens collected, however,
has "a pair of median stripes blackish pale to
pale brown, running from anterior tip of
crown to posterior margin of pronotum."
Based on the illustrations, Pseudonirvana ru-
brolimbata Kuoh & Kuoh may also prove to
be a synonym of this species.

Nirvana suturalis Melichar

Figs. 233-243

Nirvana suturalis Melichar 1903b; 166. Holotype 9, Sri
Lanka (ZMHU, examined).

Yellow. Median, longitudinal, black stripe
traversing vertex, pronotum, scutellum and
continued along inner margin of clavus of
forewing, with uneven, lateral margins and
extending angularly on corium beyond clavus
(stripe constricted a short distance from apex
of vertex in Indian specimens but not in holo-
type female) (Figs. 233, 234). First, second,
and outer margin of fourth apical cells of
forewing dark brown, costal margin with
piceous, oblique fascia about 0.25 distance
from apex; second apical cell with black spot.
Ventral surface of body creamy white.

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225 226

Figs. 221-232. Nirvana pallida Melichar: 221, head and thora.x; 222, same, profile; 223, face; 224, male pygofer; 225,
226, aedeagus, caudal and lateral views; 227, male plate; 228, connective; 229, style; 230, forewing; 231, hindwing; 232,
ovipositor.

Head narrower than pronotnm. Verte.x
about 1.5 times as long as width between
eyes, with median, longitudinal groove not
reaching apex, rugulose in anterior half, pos-
terior half polished.

Male genitalia. — Pygofer bluntly conical
caudally, with ventral, short notch, macrose-
tae confined to apical 0.33 area. Plate more
than six times as long as wide. Preapical lobe
of style rounded, relatively slender. Stem of
connective twice as long as length of each arm.
Aedeagus with short preatrium, shaft tubular,
slightly curved, each arm of dorsal apodeme
wider in middle.

Female genitalia. — Hind margin of sev-
enth sternum concave with median, short
protuberance. Ovipositor exceeding pygofer.

Measurements.— Male 4.20 (4.00-4.30)
mm long, head 0.85 (0.83-0.88) mm wide,
pronotnm 0.88 (0.85-0.90) mm wide. Female
4.74 (4.40-5.00) mm long, head 0.90 (0.80-
0.95) mm wide, pronotum 0.93 (0.88-0.98)
mm wide.

Specimens e.xamined. — Holotvpe 9 la-
beled /Type/ /7042/ /Ceylon, Nietr./ /sii-
titralis, det. Melichar/ /Nirvana suturalis
Melichar/ (ZMHU). India: 48 cJ, 66 9, col-
lected from the states of Karnataka, Kerala,

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215

Figs. 233-243. Nirvana suturalis Melichar: 233, head and thorax of holotype female; 234, female from India; 235,
face; 236, ovipositor; 237, male pygofer; 238, connective; 239, 240, aedeagus, lateral and cephalic views; 241, style; 242,
male plate; 243, forewing.

216

Great Basin Naturalist Memoirs

No. 12

and Tamil Nadu from January to December,
at an altitudinal range of 534 m (Jog Falls in
Karnataka) to 2,350 m (Ootacamund in Tamil
Nadu) above mean sea level.

Remarks. — This is the second most com-
monly collected Nirvaninae in southern In-
dia. Nirvana suturalis and N. striata are very
closely related. Both differ from other species
of Nirvana and Sophonia in having a broad,
black, median stripe running from the apex of
the vertex to the corium, but the stripe is not
as broad as in species of Chudania. Nirvana
suturalis has a serrated lateral margin to the
median stripe compared to the smoother lat-
eral margin in N. striata.

Nirvana striata, n. sp.

Figs, 244-255

Coloration as in N. suturalis, median stripe
more or less of uniform width and even mar-
gined, black spot on second apical cell of
forewing, larger than in N. suturalis, extend-
ing to first apical cell.

Head narrower than pronotum. Vertex
slightly longer than width between eyes,
anterior half sparsely rugulose. Pronotum
slightly more than 1.5 times as wide as long,
longer than scutellum.

Male genitalia. — Pygofer bluntly conical
with macrosetae confined to caudal 0.33 area,
ventral margin with short, spinelike process.
Plate caudally rounded, about seven times as
long as wide. Style with rounded preapical
lobe, apical extension of apophysis beaklike
and long. Stem of connective twice as long as
each arm. Aedeagus with shaft strongly
curved (more than in N. suturalis), apex
abruptly directed cephalad and with membra-
nous area; each arm of dorsal apodeme with
basal, short process.

Female genitalia. — Hind margin of sev-
enth sternum straight. Ovipositor exceeding
pygofer.

Measurements.— Male 4.90 (4.80-5.00)
mm long, head 0.98 (0.95-1.00) mm wide,
pronotum 1.06 (1.05-1.08) mm wide. Female
5.46 (5.30-5.60) mm long, head 1.08 (1.05-
1.10) mm wide, pronotum 1.12 (1.08-1.13)
mm wide.

Spegimens examined. — Holotype 9, In-
dia: West Bengal: Kurseong, 1,483 m,
22.x. 1981, S. Viraktamath (UAS). Paratypes:
India: Uttar Pradesh: 1 9, Mussoorie, 2,005
m, 27. iv. 1975, C. A. Viraktamath; Himachal

Pradesh: 3 9, Simla, 2,133 m, 14.x. 1979, C.
A. Viraktamath; West Bengal: 3 d, 3 9,
Kurseong, 1,483 m, 22.x. 1981, S. Virakta-
math (1 (5 , 1 9 ) and C. A. Viraktamath (2 c? , 2
9); 1 9, 15 km E of Kalimpong, 1,780 m,
27.x. 1981, S. Viraktamath; 1 <5 , 2 9 , 8 km E of
Kalimpong, 1,768 m, 29.x. 1981, C. A. Virak-
tamath (1 6 and 1 9 paratypes in BMNH,
USNM, and rest in UAS).
Remarks. — See N. suturalis.

Nirvana peculiaris, n. sp.

Figs. 256-267

Yellow. Small, black spot at apex of vertex
visible both dorsally and ventrally (absent in
female from Shillong). Fuscous, median
stripe extending posteriorly to 0.25 length
(absent in some females). Lateral, orange
stripe on vertex mesad of ocelli, continued on
pronotum, scutellum and then on to inner
claval margin where they converge and con-
tinue to apex of clavus; outer margin of stripe
on clavus often fuscous. Transverse spot at
apex of clavus, small spot at apex of inner,
anteapical cell, and anterior, oblique fascia
from costa fuscous. Black spot on each cau-
dodorsal angle of male pygofer.

Head as wide as pronotum or slightly nar-
rower. Disc of vertex depressed with rolled-
up, sharp margin, basal half with median sul-
cus, apical half with a carina, apical half of disc
very finely pitted, posterior half with minute
rugulae on polished surface.

Male genitalia. — Pygofer elongate, nar-
rowed caudally to acute, dorsally curved pro-
cess. Anal collar elongate, anterior margin
armed with two pronglike processes on each
side. Setae on pygofer confined to caudodor-
sal area where it articulates with anal tube.
Male plate elongate, 5.4 times as long as me-
dian width, macrosetae and hairlike setae con-
fined to apical half Connective robust, stem
broad, and arms as long as stem. Apophysis of
style fairly long. Aedeagus with dorsal apo-
deme poorly developed, shaft tubular, nar-
rowed caudally, strongly curved dorsally near
apex and with elongate, caudoventrally di-
rected process and two lateral, anteriorly
directed processes surrounding small gono-
pore.

Measurements.— Male 5.80 (5.70-5.90)
mm long, head 1.23 (1.22-1.25) mm wide,
pronotum 1.24 (1.22-1.25) mm wide. Female
7.40 (7.00-7.80) mm long, head 1.50 (1.45-

1988

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217

Figs. 244-255. Nirvana striata, n. sp.: 244, head and thorax; 245, face; 246, profile; 247, ovipositor; 248, forewing;
249, hindwing; 250, male pygofer; 251, male plate; 252, style; 253, 254, aedeagus, lateral and cephalic views; 255,
connective.

1.55) mm wide, pronotum 1.54 (1.50-1.60)
mm wide.

Specimens examined. — Holotype 6, In-
dia: Mizoram: Aizawl, 18. xi. 1981, C. S.
Wesley (UAS). Paratypes: India: Mizoram: 2
S,4 9, Aizawl, 18.xi.l981, C. S. Wesley; 1
d, 2 $, Lungleh, 23.xi.1981, C. S. Wesley;
Meghalaya: 1 9, Shillong, 1,961 m, 3.xi.l981,
S. Viraktamath (1 9 and 1 9 paratypes in
BMNH; 1 9 paratype in lARI, USNM, ZSI,
and rest in UAS).

Remarks. — This species is unique among
Nirvanini in having an armed tenth segment.
In coloration it resembles species of Kana,
especially K. fasciata, but it differs from those
species in having three apical cells in the hind-
wing and unique male genitalia.

Ophiuchus Distant

Ophiuchus Distant 1918b: 33. Type species: Ophiuchus
princeps Distant, by original designation and
monotypy.

Pale yellow with bright, contrasting col-
oration of black and red or pink. Head about as
wide as pronotum, spatulate, disc of vertex
depressed in middle, margined by carina, lat-
eral margin of vertex straight in front of eyes to
ocelli where it is widened and then slightly
narrowed to apex. Face with five lateral
ridges, concave medially, with short, median
carina near apex. Pronotum twice as wide as
median length, disc minutely, transversely
rugose. Hindwing with three closed apical
cells. R2 of hind tibia with 12 spines.

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No. 12

Figs. 256-267. Nirvana peculiaris, n. sp.: 256, head and thorax; 257, same, profile; 258, face; 259, female seventh
sternum; 260, forewing; 261, male pygofer; 262, male tenth segment; 263, style; 264, male plate; 265, connective; 266,
267, aedeagus, lateral and cephalic views.

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ViRAKTAMATH, WeSLEY: REVISION OF INDIAN NiRVANINAE

219

Male pygofer elongate, heavily macrose-
tose and with candal, hooklike process. Valve
semicircular. Plate rounded apically, macro-
setae scattered and confined to caudal 0.40.
Style appearing like partially closed fist.
Aedeagus large, poorly sclerotized, with ven-
tral, platelike process, shaft short, cylindrical
with large gonopore.

Remarks. — This genus appears related to
Sophonia and Nirvana with which it shares
general facies of the head. The peculiarly
shaped style and scattered macosetae on the
male plate distinguish this genus.

Ophiuchus princeps Distant
Figs. 268-276

Ophiuchus princeps Distant 1918b: 34. Lectotype 6 ,
India (BMNH, examined).

Pale yellow. Vertex laterally margined by
black line that does not reach apex, black,
median, longitudinal line does not reach apex.
Eyes centrally fuscous surrounded by red.
Ocelli red. Face ochraceous with lateral in-
foldings black. Pronotum sanguineous with
submarginal, anterior, transverse, whitish
spot. Scutellum anteriorly marked with fus-
cous band with backward, lateral projections.
Base of forewing fuscous, apical 0.33 with an
irregular, pink spot margined by fuscous, rest
yellowish.

Head shorter than combined length of pro-
notum and scutellum (36:44). Pronotum
shorter than scutellum (21:23). Face longer
than wide.

Male genitalia. — Pygofer elongate, lobe
covered with numerous stout setae and termi-
nated by curved, hooklike process. Valve
semicircular. Plate elongate, with angulate
projection on lateral margin near apex. Style
flat, preapical lobe with few setae, apophysis
with two angular projections and apically
pointed. Aedeagus with very short, dorsal
apodeme, elongate, platelike, ventral pro-
cess, armed with median ridge and two cau-
dal, spinelike processes on its laterocaudal
angles, shaft short, cylindrical, with large,
round gonopore and with pair of lateral pro-
cesses about its midlength.

Measurements. — Male 4.60 mm long,
head 1.07 mm wide, pronotum 1.05 mm
wide.

Specimen examined. — Lectotype <5 la-
beled /Travancore/ /Distant Coll. 1911-383/

/Ophiuchus princeps Dist., type/ here desig-
nated (BMNH).

Remarks. — This species has a very distinc-
tive coloration not possessed by any other
species of Nirvaninae known from the Indian
subcontinent nor by any other species of
OpJiiuchus known from the Oriental region
and Australia. The male genitalia of Ophi-
uchus species except O. distanti Evans (1973:
Fig. 5B) are not sufficiently known to suggest
any relationships among them. Judging from
the illustrations of O. distanti, it does not
appear to be related to O. princeps.

Tribe Occinirvanini

Body elongate, slightly depressed. Head
produced anteriorly, spatulate. Face horizon-
tal, frontoclypeus and clypellus either flat
{Occinirvana Evans) or convex; facial sulci
deep, prominent; face longer than wide,
genae laterally emarginate below eyes, lora
fairly prominent.

Antennae arising close to anterior margin of
head. Vertex elongate, with median sulcus
and transverse ridge or carina across ocelli,
which are situated on disc of vertex closer to
anterior than to posterior margin. Lateral
margin of pronotum rounded, without carina,
rather parallel-sided. Forewing with com-
plete and prominent venation; at least three
cross-veins reaching vein R from costal mar-
gin; anteapical cells three, outer anteapical
cell half as long as median, inner anteapical
cell open basally; apical cells four; appendix
distinct, often extending around apical margin
as in Omaranus Distant. Hindwing with three
apical cells. Hind femoral spinulation 2+1+0.
Hind basitarsus with six platellae (in Omar-
anus).

Male genitalia is of deltocephaline type.
Pygofer laterally not fused with valve, lobe
covered by stout setae. Valve triangular.
Plates triangular with stout macrosetae. Style
deltocephahne type with well-developed, pre-
apical lobe and fingerlike apophysis. Connec-
tive Y-shaped with diverging arms. Aedeagus
simple, with large gonopore, articulated with
connective.

Remarks. — The tribe, judged from the
structural features, is not closely related to
either Nirvanini or Balbillini. It has several
deltocephaline features, especially the male
genitalia.

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Great Basin Naturalist Memoirs

No. 12

Figs. 268-276. Ophiuchus princeps Distant: 268, habitus; 269, profile; 270, male pygofer; 271, male valve; 272, male
plate; 273, style; 274, aedeagus, lateral view; 275, aedeagus and connective, ventral view; 276, aedeagus, cephalic

1988

ViRAKTAMATH, WESLEY: REVISION OF INDIAN NiRVANINAE

221

Genus Omaranus Distant

Omaranus Distant 1918b: 5. Type species: Omaranus
typictis Distant, by original designation and inoiio-
typy.

Didius Distant 1918b: 36. Type species: Didius sexualis
Distant, by original designation and nionotypy.

Sexually dimorphic. Head narrower than
pronotum in male, wider than pronotum in
female. Vertex with median carina in apical
half, sulcus in basal half. Face longer than
broad, frontoclypeus tumid. Clypellus longer
than broad; frontoclypeus, lora, genae, and
clypellus with scattered, short setae. Prono-
tum gradually widened posteriorly in male,
parallel-sided and with faint, median ridge
near posterior half in female; posterior margin
strongly inwardly concave; disc llattish in fe-
male, slightly raised posteromedially in male,
lateral margins not carinate. Male forewing
exceeds abdomen, appendix runs around api-
cal margin, with four apical and three anteapi-
cal cells; hindwing well developed with three
apical cells. Female forewings cover only
three basal, visible, abdominal terga, while
hindwings cover two basal, visible terga; ap-
pendix wanting. Hind tibial spinulation Rj
16 ± 2, R, 10 ± 1, R3 3. Hind basitarsus shorter
than combined length of second and third
tarsi and terminated by six platellae.

Male pygofer without processes. Tenth seg-
ment stout, rather short. Ovipositor not ex-
ceeding pygofer.

Remarks. — This genus and the Austrahan
Occinirvana are closely related. The shape of
the head differentiates the two genera.

Omaranus typicus Distant

Figs. 277-289

Omaranus typicus Distant 1918b; 6. Lectotype S , India

(BMNH, examined).
Didius sexualis Distant 1918b: 36. Lectotype 9, India

(BMNH, examined).

Male. — Ochraceous (probably green in
life). Anterior margin of vertex narrowly black
margined, with median, black spot, two
smaller, somewhat elongate spots near poste-
rior margin, ocelli black. Dorsal-most area of
frontoclypeus and face ventrad of antennal
bases, blackish brown, marginal band dorsad
of antennal base and longitudinal stripe on
proepimeron blackish brown. Apical 0.25 of
forewing and stripe along inner margin fus-
cous. Abdomen with four black stripes; sterna
marked with brown stripes. Apical half of hind

femora ventrally streaked with brown. A ven-
tral spot near apex of fore femora black.

Genitalia. — Pygofer lobe caudoventrally
produced, caudodorsal area beset with stout,
short setae. Plate caudally rounded with scat-
tered, stout setae. Valve straplike with caudal
margin roundly and medially produced. Apo-
physis of style curved laterally with series of
teeth on lateral margin. Stem of connective
0. 75 as long as each arm . Aedeagus with short,
dorsal apodeme, shaft stout at base, narrowed
caudally in lateral aspect with a large gono-
pore on dorsal aspect.

Female. — Ochraceous. Head with median
spot on anterior margin, ocelli, and two spots
near base of vertex black; median stripe, a
transverse band across ocelli, hind margin,
except medially, and two spots in front of eyes
reddish. Dorsal margin of face black from
which two apically divergent stripes traverse
face. Proepisternum traversed by longitudi-
nal, piceous stripe, much longer on pro-
episternum; mesopleura and mesocoxae with
piceous spot. Pronotum with scattered, dark
brown spots, with obscure stripes, two lateral
and two median reddish. Scutellum except for
two lateral, basal angles in basal half black,
apical half blackish brown. Forewing veins
marked with red. Fore tibiae and tarsi red-
dish; fore- and mesothoracic femora black,
mesocoxae with piceous spot. Abdominal col-
oration as in male.

Genitalia. — Hind margin of seventh ster-
num with median, U-shaped excavation and
translucent in middle. Anal segment con-
cealed by pygofer.

Measurements. — Male 4.30 mm long,
head 1.00 mm wide, pronotum 1.10 mm
wide. Female 7.90 mm long, head 1.00 mm
wide, pronotum 1. 10 mm wide.

Specimens examined. — Lectotype 6 la-
beled / Calcutta, 28.viii.07, Mus. Coll./ /
Omaranus typicus Dist., type/ here desig-
nated. Paralectotype S labeled / Calcutta,
23.viii.07/ /Omaranus typicus Dist., type/
here designated. Paralectotype has the abdo-
men and left fore and hindwings missing. Lec-
totype 9 labeled / Calcutta/ /Distant Coll.
1911-384/ /Didius sexualis Dist. type/ here
designated (BMNH). India: Punjab: 1 c?, 1 9
nymph, Ludhiana, 13. ix. 1966, A. S. Sohi
(PAU). India: Madhya Pradesh: 3 6 , 18 9 ,
Nebudda Survey, Sta. No. 31, 37, 58,
21.ii-9.iii.l927, H.S. Pruthi (ZSI).

222

Great Basin Naturalist Memoirs

No. 12

Figs. 277-289. Omamnus ti/picus Distant: 277, head and thorax of lectotype male; 278, same of lectotype female,
Didius sexiialis Distant; 279, ovipositor; 280, 281, fore and hindwings of male; 282, male pygofer; 283, male plate; 284,
male valve; 285, connective; 286, style; 287, 288, aedeagus, lateral and cephalic views.

1988

ViRAKTAMATH, WeSLEY: REVISION OF INDIAN NiRVANINAE

223

Acknowledgments

The authors express thanks to Dr. W. J.
Knight, Mr. M. D. Webb (BMNH), Dr. U.
Gollner-Scheiding (ZMHU), Dr. H. Synave
(IRSNB), Dr. A. S. Sohi (PAU), and Dr.
Baldev Sharma (JU) for lending specimens for
study, including primary types under their
care. Also, the director of the Zoological Sur-
vey of India allowed the senior author to ex-
amine Dr. H. S. Pruthi's type material during
his visit to ZSI. The authors are also grateful to
Dr. P. H. Freytag, University of Kentucky,
Lexington, for critically reviewing the manu-
script.

Literature Cited

The references cited in the text prior to 1956 are found
in Metcalf (1964), wherein Pruthi s works have been cited
under the name Singh-Pruthi.

Ahmed, M., and S. H Mahmood 1970. A new genus and
two new species of Nirvaninae (CicadeUidae: Ho-
moptera) from Pakistan. Pakistan Journal of Scien-
tific and Industrial Research 12: 260-263.

Ahmed, M , and K F Mallik 1972. On the true identifi-
cation of genus Chudania Distant collected on fig
{Ficiis carica ) in forests of Azad Kashmir and Kap-
tai (Pakistan). Pakistan Journal of Forestry 22;
103-108.

Davis, R. B. 1975. Classification of selected higher cate-
gories of Auchenorrhynchous homoptera (Cica-

deUidae and Actalionidae). Technical Bulletin
No. 1494, Agricultural Research Service, United
States Department of Agriculture. .52 pp.

EvAN.s. J W 1966. The lealhoppers and froghoppers of
Australia and New Zealand (Homoptera: Cicadel-
loidea and Cercopoidea). The Australian Museum
Memoir 12: 1-347.

1973. Some new genera and species of Cicadel-

loidea from Australia and New Guinea (Ho-
moptera). Pacific Insects 15: 185-197.

KUOH. C L , AND J L KUOH 1983. New species of
Pseudonirvana (Homoptera: Nirvanidae). Acta
Entomologica Sinica 26: 316-.325 [In Chinese].

Kramer, J P 1964. A review of the Neotropical Nirvan-
inae (Homoptera: CicadeUidae). Entomological
News 75: 113-127.

LiNNAVUORl, R 1979. Revision ofthe African CicadeUidae
(Homoptera Auchenorrhyncha), Part II. Revue de
Zoologie Africaine 93: 929-1010.

Metcalf, Z P 1963. General Catalogue of the Ho-
moptera, Fascicle VI. Cicadelloidea, Part 7 Nir-
vanidae. Agricultural Research Service, United
States Department of Agriculture. 35 pp.

1964. General Catalogue of Homoptera, Fa.scicle

VI. Cicadelloidea, Bibliography of the Cicadel-
loidea (Homoptera: Auchenorrhyncha). Agricul-
tural Research Service, United States Depart-
ment of Agriculture., 349 pp.

Ramakrishna, B V. 1980. Leaihopper fauna of pulse
crops and biology of Empoasca (Distantasca) ter-
minalis Distant (Homoptera: CicadeUidae). Un-
published thesis. University of Agricultural Sci-
ences, Bangalore. 197 pp.

Theron, J G 1970. A new species of Narecho
(Hemiptera, Cicadelloidea: Nirvanidae) from
Africa. Novos Taxa Entomologicos No. 83, 1-7.

GENUS FLEXAMIA: NEW SPECIES, PHYLOGENY, AND ECOLOGY

Robert F. Whitcomb' and Andrew L. Hicks'

Abstract — Eight new species of Flexamia DeLong are described, and two Mexican species previously syn-
onymized are reinstated. Species are assigned to 13 species groups; keys are presented for the groups and for the 44
recognized species. Host data, in many cases with ohgophagy coefficients, are presented for 37 of the species. Many
species speciaUze on native, dominant, perennial, choridoid or panicoid grasses; some are monophagous. Seven sister
species pairs specialize on the same (or a closely related) grass species; in addition, four closely related species appear to
be restricted to Bouteloua curtipendtila. Few if any specialists colonize their host throughout its entire range.
Ecological factors such as phenology and/or host patchiness strongly influence geographic distribution. An intuitive
phylogeny is proposed and is used as a basis for constructing 18 character transformations. The genus Flexamia
probably originated in Mexico by division of an ancestral lineage from which the modern genus Spartopyge also
diverged.

The first specimen of the genus Flexa7nia
DeLong was apparently collected at Jack-
sonville, Florida, in the height of the Lin-
naean explosion; it was deposited in the
British Museum (Natural History) and named
Jassus productus by Walker (1851). In their
studies of Iowa prairies in the latter part of the
nineteenth century, Osborn and Ball encoun-
tered other species, which they referred to
the genus Deltocephalus. DeLong (1926)
erected a subgenus Flexamia DeLong within
Deltocephalus Burmeister. DeLong and
Sleesman (1929) recognized that Delto-
cephalus was an unworkably large assemblage
of distantly related species and divided the
genus into several genera (e.g., Laevi-
cephalus DeLong & Sleesman, Pohjamia De-
Long & Sleesman, and Flexamia). The genus
concept for Flexamia was refined by Oman
(1949) in his revision of North American ci-
cadellid genera, but the modern generic con-
cept was established by Young and Beirne
(1958). These authors segregated Spartopyge
Young & Beirne, synonymized Secopennis
Oman, described genitalic characters of males
and females (the first use of genitalic charac-
ters for Flexamia females), and presented a
species key. The 30 species they described
included all of the widely distributed and
commonly collected representatives of the
genus, and their key proved to be accurate
and easy to use. Young and Beirne also de-
scribed possible relationships among species

and, without specifying an explicit phylogeny,
proposed broad outlines of hypothetical evo-
lutionary events in the genus. The genus was
incorrectly synonymized with Acurhinus Os-
born by Linnavuori (1959). Comparison of
the specimens (all females) of Acurhinus
( =Dorydium) maculatum Osborn in the Ohio
State University collection led Linnavuori
and DeLong (1978) to synonymize Acurhinus
with the African stirelline genus Hododoecus
Jacoli. The genus Flexamia as defined herein
is an entirely North American taxon related to
Spartopyge, to Alapus DeLong & Sleesman,
and, perhaps distantly, to Aflexia Oman. The
closest Old World relative may be Enantio-
cephalus Haupt.

Although the taxonomic status of Flexamia
has been generally adequate since 1958, host
data have been difficult to obtain. Earlier col-
lectors were not knowledgeable agrostologists
and often reported only that deltocephaline
species, including Flexamia, resided on
"grasses. " Astute collectors such as Osborn
and Ball recognized the importance of biologi-
cal data, and their notes contain some host
records. In 1967, H. H. Ross encouraged the
senior author to ascertain the hosts of Flex-
amia species, in large part to test his intuitive
belief that host relationships, especially host
transfers, could be invoked to explain specia-
tion of phytophagous insects. Accordingly, we
sought to determine hosts of the species de-
scribed by Young and Beirne. Much to our

'insect Pathology' Laboratory, AEriciiltural Research Service, Beltsville, Maryland 20705.

224

1988

Whitcomb, Hicks: Flexamia Revision

225

Fig. 1. Flexamia serrata.

surprise, in the course of this search, unde-
scribed species emerged, particularly when
we examined dominant, perennial Muhlen-
bergia or Bouteloua species (Whitcomb et al.
1986, 1987). Under these circumstances, it
seems appropriate to review the species con-
cept and to emend the generic concept,
which, although basically intact in its essential
aspects, must accommodate the new species.

Genus Flexamia DeLong

Deltocephalus subgenus Flexamia DeLong 1926:20, 22.

Type species: Deltocephalus reflextis Osborn &

Ball 1897, by original designation.
Flexatnius, DeLong and Sleesman 1929:82.
Flexamius subgenus Secopennis DeLong & Sleesman

1929:85. Type species: Deltocephalus slossoni

Ball.
Acurhinus Oshorn, Linnavuori 1959:89.
Flexamia DeLong, Young and Beirne 1958; Linnavuori

and DeLong 1978:208.

Small deltocephaline leafhoppers. Length
of male 2.4-5.0 mm, of female 2.6-5.0 mm.
Head produced, median length of crown usu-
ally greater than interocular width; clypellus
broader at base than at apex; forewing with
reflexed veinlets to costal margin in vicinity of
small, outer anteapical cell.

Dried specimens yellowish brown or gray,
with ferruginous to black markings; live in-
sects in some cases with green pigmentation
changing to orange or stramineous when
dried. Apical area of crown partly or com-
pletely surrounded by darker markings, in
latter case with clearly defined, apical areolet.
Crown often with pair of oblique, dark mark-
ings at base; pair of transverse, dark markings
at about midlength; pair of darker markings
parallel to lateral margins near apex. Face
color varying from pale to black, often useful

in species discrimination. Pronotum of some
species with conspicuous, longitudinal
stripes, but many species with faint, wider
stripes. Veins of forewings usually dark-mar-
gined; distinct, dark-margined areole fre-
quently near center of corium.

Male, — Plates usually shorter than pygo-
fer, frequently fused along mesal margins;
pygofer usually with pronounced posterior
lobe in lateral aspect; style with longitudinally
grooved, preapical lobe, which is usually not
pronounced; connective linear, arms contigu-
ous basally, sometimes fused to aedeagus.
Connective with upper portion less heavily
sclerotized than lower portion, often thinner
and extended dorsad in thin keel that varies in
width interspecifically; dorsal aedeagal apo-
deme with pair of conspicuous appendages
directed caudoventrad or caudad. Aedeagus
interspecifically variable, of great value in
species discrimination.

Female. — Sternum VII nearly always
longer than VI, often 2X or more; hind margin
with 2-4 teeth and usually small, median
notch. Pygofer with well-dispersed setae;
ovipositor barely reaching pygofer apex.

Flexamia Species Groups

In the following discussion, we describe 13
species groups and provide keys to species.
We describe 8 new species and reinstate 2
species previously synonymized. We also re-
view important characteristics of previously
described species, emphasizing diagnostic
morphology, ecology, and biogeography. Our
brief descriptions of previously described spe-
cies complement the descriptions of Young
and Beirne (1958).

Key to Flexamia Species Groups

1 . Dorsum with pair of stripes extending from hind
portion of disc of crown across pronotum and
scutellum (Figs. 2A-D) 2

— Dorsum without such stripes (Figs. 2E,F) 4

2(1). Individuals large, at least 4.5 mm

IV. grammica group

— Individuals smaller 3

3(2). Forewing broadly truncate (Fig. 2D); Arizona

VIII. ritana group

— Forewing obliquely truncate (Fig. 2B) or not

truncate (Fig. 2A) (prairie); Florida

VII. alhida group

4(1). Face entirely black, genae at most with oblique,
pale streak; crown without midlength transverse
lines 5

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Great Basin Naturalist Memoirs

No. 12

Fig. 2. Habitus ofFlexamia species: A, alhida; B, shssonae; C, grammica; D, ritana; E, pijrops: F, areolata.

1988

Whitcomb, Hicks: Flexamia Revision

227

Fig. 3. Faces of Flexamia species: A, serrata- B, stylata; C, prairiana- D, prairiana; E, picta; F, decora; G,flexulosa;
H, sandersi.

— Face black basally, pale apically, or brownish
in part; at least genae pale, crown usually with
midlength transverse lines 6

5(4). Forewing with conspicuous black spot (Fig.
2F) in corium; crown with median length
more than one-half greater than interocular
width XII. areolata group

— Forewing without conspicuous black spot in
corium; crown with median length less than

one-half greater than interocular width

XI. impiitans group

6(4). Face pale with several narrow, parallel, fus-
cous, interocular lines interrupted at middle

(Fig. 3A); northwestern (Fig. 30)

XI. serrata group

— Face with (Figs. 3B,D-G) or without (Figs.
3C,H) interocular lines but not as above 7

7(6). Either with crown more than twice interocu-
lar width (Fig. 2E) {pyrops Crumb), or with
conspicuous, dark, apical wing blotch (Fig.
4C) and sordid yellow face with black interoc-
ular band {picta [Osborn]); viewed laterally,
male pygofer with conspicuous, acutely angu-
late, ventral lobe (Figs. 5A,B); aedeagus (Fig.
6D) with pair of anteapical processes and api-
cal, flangelike processes VI. picta group

Habitus not as above; male pygofer with pos-
terior lobe rounded ventrally; aedeagus not as
above 8

8(7). Males 9

— Females 14

9(8). Aedeagus and connective fused (Figs. 6F,G)

XIII. prairiana group

— Aedeagus and connective distinctlv articu-
lated (Figs. 6A-C, E) ' 10

10(9). Aedeagus without apical processes (Fig. 6C) 11

— Aedeagus with (at least minute) apical

processes 12

11(10). Male plates broad, not tapered (Figs. IIA-I)

I. pectinata group

— Male plates more elongate and tapered (Figs.
9C,F) III. zacate group

12(10). Aedeagus (Figs. 7F,G) with 2 pairs of pro-
cesses V. curvata group

— Aedeagus not so 13

13(12). Aedeagus with 2 processes on shaft (Fig. 7H)

II. abbreviata group

— Aedeagus with 3 processes on shaft (Figs. 6E,
31A,D-M) X. flextdosa group

14(8). Face white, ivory, or very pale yellow,
sharply contrasting with black interocular line
(Figs. 3B,F,G) X. flexulosa group (in part)

— Face with lower part not white or pale (Fig.
3y), or if pale, not contrasting sharply with
interocular band, or interocular band brown,
not black (Fig. 3D) 15

15(14). Body color stramineous 16

— Body color not stramineous 17

16(15). Hind margin of sternum VII with medial pro-
jection (Fig. lOO)

X. flexulosa group (inflata subgroup)

— Hind margin of sternum VII without projec-
tion 20

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Great Basin Naturalist Memoirs

No. 12

Fig. 4. Wings of Flexamia species: A, inflata; B, cur-
vata; C, picta.

17(15). Sternum VII (Fig. lOE) with median, trape-
zoidal projection, with a medial notch, usu-
ally with smaller, shallower notches on either
side II. abbreviata group

— Sternum VII not as above 18

18(17). Sternum VII produced medially in a distinct

lobe or projection (Figs. 10S,T)

XIII. prairiana group

— Sternum VII not produced medially in dis-
tinct lobe, or if produced, with deep, median
incision 19

19(18). Hind margin of sternum VII incised to form
pattern of four teeth; hind margin usually
heavily infuscated on hind margin in vicinity
of teeth (Figs. 14A-H) I. pectinata group

— Hind margin of sternum VII not so 20

20(19). Hind margin of sternum VII slightly concave
on each side of median excision; infuscated
notches usually contrasting sharply with
stramineous color of sternum (Figs. 10J,K)
V. curvata group

— Hind margin of sternum VII not so (Figs.
101, L) III. zacate group

I. The pectinata Group
The greatest proposed changes in Flexamia

species concepts are in the pectinata species
group to which we add five new species and
reinstate two others. All species of the group
have aedeagal shafts without apical processes.
Although aedeagal characters are not as useful
for species discrimination in this group as in
other groups, the male plates and female ster-
num VII provide useful apomorphies that de-
fine the group. The plates (except in collo-
rurn) are broad and basally parallel-sided. The
hind margins of the female sternum VII of all
species possess four medial teeth with sur-
rounding infuscation (but note the extensive
modification in mescalero [Fig. 21H]). The
size and shape of the teeth vary interspecifi-
cally. The species are largely southwestern
and/or Mexican, and many appear to have
small ranges. The nominate species F. pecti-
nata, however, has a wide geographic range.
As defined by Young and Beirne (1958), this
species ranges from western Mexico {zamora
DeLong & Hershberger) through the eastern
highlands (Valles and Monterrey [7ninima
DeLong & Hershberger]), the Davis Moun-
tains of west Texas, Otero and Guadalupe
counties of New Mexico north to North Da-
kota, and as far east as Akron, Ohio, in the
prairie peninsula. However, our studies indi-
cate that zamora and minima should be rein-
stated. Recognition herein of the new species
batidarita, gila, coUorum, jacala, and mesca-
lero makes it clear that this group is diverse
and has speciated frequently. The host of
pectinata is side-oats grama (Bonteloua cur-
tipendula), a widespread prairie species. In
addition to pectinata, there are three south-
western side-oats grama specialists of the
pectinata group. Flexamia doeringae and F.
gila reside in the southeastern Arizona moun-
tains and Mogollon Rim region and extended
conifer-oak savanna of Mexico, respectively.
Flexamia bandarita, as far as known, is con-
fined to the Chisos Mountains and Marathon
Basin of Trans-Pecos Texas. We suspect that
side-oats grama, which also has a wide range
in Mexico, will prove to be the host of some
Mexican species of the pectinata group. The
most distinctive species of the group {mesca-
lero), however, appears to specialize on Muh-
lenhergia pauciflora.

Several factors may account for speciation
in the pectinata group. Side-oats grama is it-
self a complex of ecotypes that vary greatly in
structure and seasonality (Gould 1979). This

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Whitcomb, Hicks: Flexamia Revision

229

Fig. 5. Pygofers oi Flexamia species: A, picta; B, pyrops; C, stylata; D, producta. Redrawn from Young and Beirne

(1958).

Fig. 6. Aedeagus and connective of Flexamia species: A, curvata; B, abbreviata; C, canyonensis; D, picta; E,
flexulosa; F, prairiana; G, graminea. Redrawn from Young and Beirne (1958). Aspects; A-F, caudoventral; G, lateral.

chloridoid grass crosses many biome lines and
is a dominant in mesic Sonoran and Chi-
huahuan grasslands and in montane systems
within these semiarid grasslands. It is com-
mon through much of the prairie and occurs
eastward on nonforested slopes in the Ap-
palachian mountains. Like other major domi-
nants of North American grasslands, there-
fore, side-oats grama has a much wider
geographic range than any of the cicadellid
species that exploit it.

The Chihuahuan desert system presents a
major barrier to geographical continuity of
prairie grasses, and side-oats grama is no ex-
ception. Many prairie cicadellid taxa occur
west of this desert system, but, depending
upon the case, the taxonomic results vary. For
example, populations related to the buffalo-
grass specialist Athysanella (Gladionura)
emarginata occur in disjunct stands of
Buchloe in New Mexico's Hidalgo County.
This population, until very recently, had been
regarded as a subspecies (sobrina) but will be
elevated to a species in a forthcoming revision

oi Gladionura (H. D. Blocker, personal com-
munication). On the other hand, forms of A.
(Gladionura) argenteola (a Botiteloua gracilis
specialist) in southeastern Arizona are not dis-
tinguishable from short-grass prairie forms.
The connecting link between the relatively
mesic, temperate grasslands of the northern
Mexican highlands and the prairie is tenuous
at best, consisting of a small set of "stepping
stone" montane islands within the Chi-
huahuan desert of Coahuila and Chihuahua,
or of the relatively narrow grassland periph-
ery surrounding the desert on the east and
west and meeting at the south in the state of
San Luis Potosi. Also, the mesquite-acacia
savanna and Trans-Pecos shrub savanna of
Texas are further barriers to geographic conti-
nuity of B. curtipendula. During full glacia-
tions, these arid regions had climates that
were much more moist than they are today
(Van Devender and Burgess 1985).

One of the new species, collorum, from the
Edwards Plateau of central Texas, is appar-
ently a specialist on Nealley grama, Bouteloua

230

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LT CA

CV LT

Fig. 7. Aedeagal apices oi Flexamia species; A, scrrata- B, alhida; C, slossonac- D, litana- E, ^rammica; F, curvata;
G, sitrcuhi; H, abbreviatii; I, zflcafc, J, canijonensis; K, /;icf«, L, ptjrop.s. Aspects: AD, anterodorsal; CA, caudal; CV,
caudoventral; DS, dorsal; LT, lateral. Redrawn from Young and Beirne (1958).

uniflora. Flexamia coUonim may be related to
another new species, jacala, whose type lo-
cahty in Mexico is close to the intersection of
eastern and western montane Mexican grass-
lands. The latter species is represented by a
single male specimen from the USNM, so
nothing is known about its biology.

Finally, another new species, niescalero,
discovered in a single location in southeastern
New Mexico, presumably arose by host trans-
fer to New Mexican muhly, Muhlenbergia
paucijlora.

Description of the pectinata Group

Medium-sized to small deltocephaline leaf-
hoppers. Length of 6 2.7-3.8 mm, of 9
2.8-4.2 mm. Species with general facies of
genus; brown or in dark specimens, almost
black. Crown variably produced. \6] Plates

(Figs. IIA-I) elongate, broad, parallel-sided
basally (except coUorum). Pygofer with dis-
tinct, posterior lobe, strongly constricted in
lateral aspect, both dorsally and ventrally.
Aedeagus and connective distinctly articu-
lated. Aedeagus (Figs. 12A-I) with no apical
processes, or at most slightly capitate
(mescalero), often appearing flared in dorsal
aspect. [Note: Caution must be exercised
when using the aedeagal apex as a diagnostic
character, since this structure is membranous
and is subject to tearing and breaking during
copulation or preparation of the specimen.]
Styles variable interspecifically (Figs. 13A-I).
Gonopore subapical on caudoventral surface.
[ 9 ] Sternum VII (Figs. 14A-H) with four me-
dial teeth on hind margin; teeth varying inter-
specifically in length and shape, surrounded
by infuscated region.

1988

Whitcomb, Hicks; Flexamia Revision

231

Fig. 8. Male plates and pygolers oi Flexamia species: A, alhida: B, slossonae: C, serrata, D,ritana; E, abbreviata; F,
grammica.

Fig. 9. Male plates and pygofers oi Flexamia species: A, picta; B, pyrops; C, canyunensis; D, curvata;
E, surcula; F, zacate.

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No. 12

Fig. 10. Female sternum VII of Flexarnia species: A, albida^ B. slossonac, C, ^errata^D, t"7;,L" o'llXa- P
grammica; G, picta; H, pyrops; I, canyonensis;], surcula; K, curvata; L, zacate; M,flexulosa; N, decora, O, mjlata, F,
imputans; Q, areolata; R, sfyiata; S, prainana; T, sandersi; U, fitammea.

1988

Whitcomb, Hicks: Flexamia Revision

233

Fig. 11. Male plates and pygofers oiFlexamia pectinata group, ventral aspect: A, mminui; B
D, handarita- E, gila; F, doeringae; G, collorurn; HJacala; I, mescalero.

zamora; C, pectinata;

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No. 12

A B u u t . - ^

Fig. 12. Aedeagal apices of Flexamia pectinata group, caudoventral aspect: A, minmui; B, zainora; C, pectinata; D,
bandarita; E, gila; F, doeringae; G, coUorum; H,jacala; I, mescalero.

Key to Males of the pectinata Group
1. Plates long and parallel-sided at bases 2

— Plates short, not parallel-sided

collorum Whitcomb & Hicks

2(1). Plates subrectangular (Figs. IIC.F) 3

— Plates with apices rounded 4

3(2). Aedeagus in ventral aspect (Fig. 12F) with pro-
tuberance close to tip; face pale; Arizona

doeringae Beamer & Tuthill

— Aedeagus in ventral aspect with protuberance
considerably removed from tip (Fig. 12C); face
brown; prairie pectinata (Osborn & Ball)

4(2). Plates (Fig. 11 A) with apices acute; Mexico . . .

minima DeLong & Hershberger

— Plates with apices not acute 5

5(4). Plates almost as long as pygofer (Fig. HE) ....

gila, n. sp.

— Plates no more than 3/4 length of pygofer 6

6(5). Aedeagus expanded apically in ventral aspect,
appearing bluntly saggitate (Figs. 121, 22) ....
mescalero, n. sp.

— Aedeagus not expanded apically 7

7(6). Style apex (Fig. 13H) avicephaliform in lateral

aspect jacala, n. sp.

— Style apex not avicephaliform 8

8(7). Style apex (Fig. 13D) ventrally directed, chelate

bandarita, n. sp.

— Style apex (Fig. 13B) not chelate

zamora DeLong & Hershberger

Key to Females of the pectinata Group

1. Sternum VII with 4 teeth of approximately equal

length and 3 relatively shallow incisions 2

— Sternum VII with deep incisions and/or teeth of
different length 4

2(1). Sternum VII with infuscations on either side of
middle incision (Fig. 14G); Texas, host Butite-
loua uniflora coUorum, n. sp.

— Sternum VII not as above 3

3(2). Sternum VII with heavy infuscation around

teeth on hind margin (Fig. 14C); USA, prairie
pectinata (Osborn & Ball)

— Sternum VII with light infuscation on hind mar-
gin (Fig. 14A); Mexico

minima DeLong & Hershberger

4(1). Sternum VII with outer teeth longer than inner

teeth 5

— Sternum VII with inner teeth longer than outer
teeth 6

5(4). Sternum VII with outer teeth sharply pointed,
median incision shallow (Fig. 14B); Mexico . . .
zamora DeLong & Hershberger

— Sternum VII with outer teeth blunt, median
incision deep (Fig. 14H) mescalero, n. sp.

6(4). Inner teeth of sternum VII together forming

blunt, median projection (Fig. 14F)

doeringae Beamer &"Tuthill

— Inner teeth of sternum VII separated, acute at
tip, separated by tapered incision 7

7(6). Sternum VII with three incisions of equal depth

(Fig. 14E) gila, n. sp.

— Sternum VII with median incision deeper than
lateral incisions (Fig. 14D) bandarita, n. sp.

1. Flexamia minima DeLong &
Hershberger, n. stat.

Flexamia minima DeLong & Hershberger 1947: 138.
Flexamia pectinata. Young & Beirne 1958: 46. (In part.)

Length of c? 2.9 (2.8-2.9) mm, 9 3.0
(2.7-3.2) mm; head width of 6 0.84 mm, 9
0.92 mm. Crown produced; median length of
crown 0.74 x head width and L45 x interocu-
lar width (c? n = 6; 9 n = 3). Crown hght
brown to pale yellow with dark, circular spot
around apex, pair of broken, transverse lines
at midlength, two obli(|ue markings at rear,
appearing continuous with the medial pair of
six pale brown, pronotal stripes. Face with
broad, dark brown, interocular band, fading
to pale yellowish. Brown basally, disc and

1988

Whitcomb, Hicks: Flexamia Rf.vision

235

blX

Fig. 13. Styles and style tips oiFlexamia pectinata group: A, minima; B, zamora; C, pectinate; D, bandarita; E, gila;
F, doeringae; G, collorum; H,jacala; I, mescalero.

apex of forewings usually with irregular, dark
spot.

Male. — Pygofer much like that o( pectinata
but shorter; plates (Fig. 11 A) barely exceed-
ing ventral lobe in lateral view or 2/3 length of
pygofer, unevenly tapering to blunt apices.
Genitalia similar to those of pectinata and
zamora, but little more than half their size. In
ventral aspect, apex of style not exceeding
plane of well-developed, preapical lobe, angle
formed between preapical lobe and style apex
more acute than in pectinata. Connective
similar to that of pectinata, but with a nar-
rower, deeper, U-shaped incision at the joint
with the aedeagus. Other characters as in
pectinata.

Female. — Sternum VII with posterior
margin broadly, shallowly emarginate, hind
margin with four teeth of approximately equal
length (Fig. 14A), lightly infuscated. Oviposi-
tor with bases of first valvulae as in zamora.

Types. — Holotype i: Valles, San Luis Po-
tosi, Mexico, 1 December 1938, J. S. Cald-
well.

Remarks. — An additional series o{ minima,
collected by E. D. Ball at Monterrey, Nuevo
Leon (5d, 39, 11 August 1936, USNM), has

been examined. On the basis of characters of
the male plates, the styles, and the distinc-
tiveness of the female sternum VII from that
of zamora, we reinstate minima. Species
problems in this group are discussed under
"Species Concept." This species, like zamora,
may be entirely Mexican in distribution and is
represented by the two collections discussed
above (Fig. 15). The host or hosts are un-
known, but the closely related pectinata spe-
cializes on Bouteloua curtipendula. The
grasslands of San Luis Potosi are extensive,
and many other warm season grasses, includ-
ing other Bouteloua spp., are present (Rze-
dowski 1966); further fieldwork is required to
clarify the biology oiininima.

2. Flexamia zamora DeLong &
Hershberger, n. stat.

Flexamin zamora DeLong and Hershberger 1947: 137.
Flexamia pectinata. Young and Beirne 1958: 46. (In part.)

Length of c? 3.6 (3.5-3.7) mm, 9 3.9
(3.7-4.2) mm; head width of S 1.08 mm, 9
1.15 mm. Crown not strongly produced; me-
dian length of crown 0.61 x head width and
1.27 X interocular width ((5 n = 7; 9 n = 3).

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Great Basin Naturalist Memoirs

No. 12

Fig. 14. Female sternum VII ofFlexamia pectinata group: A, minima: B, zamora: C, pectinata: D, bandarita; E, gila;
", doeringae; G, collorum; H, mescalero.

Crown light brown with dark, circular spot
around apex, pale brown, transverse line on
either side at middle. Pronotum light brown,
usually with traces of four wide longitudinal
stripes. Elytra subhyaline, dark brown spot
on disc; costal veinlets heavily margined with
dark brown. Face black above, pale brown on
lower portion.

Male. — Pygofer as in pectinata. Plates
(Fig. IIB) extending to approximately 3/4
length of pygofer, with apices less produced
than in pectinata, bluntly rounded. Style
apices exceeding the plane of the moderately
developed, preapical lobe, forming an angle
with it smaller than that in pectinata. Other
characters as in pectinata.

Female. — Sternum VII (Fig. 14B) with
posterior margin shallowly emarginate; four
produced teeth at middle; outer teeth longer,
acute, curved inward. Ovipositor with basal
processes of first valvulae similar to those of
pectinata, but recurved portion not separated
along the middle.

Types. — Holotype 6: Zamora, Michoacan,
Mexico, 2 October 1941, Plummer, Good,
Caldwell and DeLong (OSU). Paratvpes: 5 6 ,
3 $ , same collection data (OSU and USNM).

Remarks. — Although the male genitalia of

zamora are similar to those o{ pectinata, the
styles oi zamora differ in having a more acute
angle between the apex and the less well-
developed preapical lobe. Also, the plates are
more evenly rounded than those of pectinata
and longer than those of minima. The female
sternum VII of zamora differs from that of
pectinata in that the outer teeth of the hind
margin are much longer than the inner teeth;
in pectinata and minima the teeth are of equal
length. This species, almost certainly entirely
Mexican in distribution, is known only from
its type locality (Fig. 15). The host or hosts are
unknown, but other members of the complex
feed on Bouteloua curtipendula.

3. Flexamia pectinata (Osborn & Ball)

Dehocephahis pectinatus Osborn & Ball 1897: 205.
Deltucephalus (Flexamia) pectinatus DeLong 1926: 32.
Flexamius pectinatus. DeLong and Sleesman 1929: 83.
Flexamius zatnora. Young and Beirne 1958: 46. Incorrect

synonymy.
Flexamia minima, Yoimg and Beirne 19.58: 46. Incorrect

synonymy.

Important CHARACTERS. — Length of S 3.4
(2.9-3.8), of 9 3.6 (3.1-4.0). Head slightly
produced (median crown length 1.40 x inter-
ocular width; 0.62 X head width) (c? n = 37;

1988

WHITCOMB, HiCKS: FLEXAMIA REVISION

237

Fig. 15. Geographic distribution of Flexaniia pectinata {i
miniirui (O), And jacala (ir) with Botiteloua curfipendtila.

), doeringae (■), bandarita (A), gila (D), zatnora (A),

9 n = 29). [6] Pygofer (Fig. 62F) strongly
constricted both ventrally and dorsally; plates
(Fig. IIC) wide and parallel-sided for much of
their length, almost subrectangular, but with
rounded tips. Aedeagus (Fig. 12C) symmetri-
cal, without lateral processes; gonopore sub-
apical on caudoventral surface. [9] Sternum
VII (Fig. 14C) with four teeth of approxi-
mately equal length. Ovipositor with bases of
first valvulae recurved and calipterate (Fig.
63G).

Geographic distribution. — This species
is a specialist of side-oats grama {Bouteloiia
curtipendula) in prairie and mesic grasslands;
it occurs (Fig. 15) at higher elevations of
the desert plains of eastern New Mexico east
to Akron, Ohio, the eastern extension of the

prairie peninsula (Transeau 1935).

Biology. — Like other Flexamia species,
pectinata is probably multivoltine and is
present throughout the growing season. In
New Mexico it is often absent from its host
(Botiteloua curtipendula, side-oats grama) at
lower elevations where host growth may be
unpredictable because of sporadic drought. In
Chihuahuan and Sonoran highlands it is re-
placed by bandarita and doeringae, respec-
tively, and in the intervening mountains, by
gila.

Oligophagy coefficients. — Gramineae
1.000; Chloridoideae 0.973; Bouteloua 0.926;
Bouteloua curtipendula 0.825 (n = 57).

Remarks and diagnosis. — Flexamia pecti-
nata can be recognized by a combination of its

238

Great Basin Naturalist Memoirs

No. 12

Fig. 16. Flexamia gila, n. sp. : A, aedeagus and connective, lateral aspect; B, aedeagus and connective, dorsal aspect;
C, apex of aedeagus, caudoventral aspect; D, male plates and pygofers, ventral aspect; E, male pygofer, lateral aspect;
F, right style, dorsal aspect; G, ape.x of right style, lateral aspect; H, female sternum VII; I, bases of first valvulae of
female, dorsal aspect.

external genitalic characters and its geo-
graphic range. Other species (doeringae, ja-
cala) of the pectinata group with somewhat
similar male plates do not occur with pecti-
nata in its range. Of the nine members of the
group, only mescalero (host: Muhlenhergia
pauciflora) and colloruin (host: Bouteloua
uniflora ) have ranges that overlap the range of
pectinata. The male plates (Fig. Ill) and fe-
male sternum VII (Fig. 14H) of mescalero
distinguish it from pec^inaf a (Figs. IIC, 14C).
The male plates oi collorum (Fig. IIG) differ
from those of pectinata in general aspect and
in the absence of the sclerotized ridge on the
inner plate surface that characterizes coUo-
rum. The female sternum VII of collorum is
similar to that ofpectinata. Although the char-
acters discussed above permit tentative
recognition of the commonly encountered
pectinata, definitive identification requires
examination of the genitalia.

4. Flexamia gila, n. sp.
Length of 6 3.4 (3.1-3.7) mm,

9 3.8

(3.6-4.0) mm; head width of d 1.03 mm,
9 1.08 mm. Crown not strongly produced;
median length of crown approximately
0. 63 x head width and 1 . 32 x interocular width
(d n = 6; 9 n = 4).

Crown light brown with dark, circular spot
around apex, pale brown, transverse line on
either side at middle. Pronotum light brown,
usually with traces of four wide, longitudinal
stripes. Elytra subhyaline, dark brown spot
on disc; costal veinlets heavily margined with
dark brown. Face black above, pale brown on
lower portion.

Male. — Pygofer (Fig. 16E) much as in ban-
darita; posterior margin slightly more pro-
duced. Plates (Fig. 16D) broad, gradually
rounded from base to apex, extending to ap-
proximately 4/5 length of pygofer. Connective
in lateral view with dorsal keels narrow (or
rarely absent), approximately 1/4 height of
dorsal apodeme; apodemal processes as in
bandarita; styles (Fig. 16F) long, diverging
apically in ventral aspect, preapical lobe
rounded, produced ventrad and laterad; style

1988

Whitcomb. Hicks; Flexamia Revision

239

portion distad from preapical lobe gradually
narrowing to embrowned, strongly divergent,
acute apex in ventral aspect, or, in lateral
aspect (Fig. 16G), appearing bladelike, pro-
duced ventrally and flattened, bearing minute
teeth along the irregularly curved ventral
margin of style ending in a truncate apex that
is slightly curved ventrad. Aedeagus (Figs.
16B,C) symmetrical, short, slender. Apical
margin flared and convex, much as in pecti-
nata. Distance between gonopore and apex
less than in pectinata. Gonopore subapical on
caudoventral surface.

Female.— Sternum VII (Fig. 16H) with
posterior margin broadly but shallowly emar-
ginate, with four produced teeth; middle
teeth broad. Ovipositor with basal processes
of first valvulae (Fig. 161) recurved; recurved
portion constricted above proximal margin,
expanded slightly mesad and distad into dif-
ferentially sclerotized lobes.

Types. — Holotype 6 : New Mexico, Sierra
Co., Kingston, 17 August 1985, R. F. Whit-
comb (6,300 ft, IPL 001875, Bouteloua cur-
tipendula). Paratypes: 3 6 and 1 9, same
collection data; 2 6 and 2 9 , Mexico, 5 km S
Casas Grandes, Chihuahua (5,600 ft, Boute-
loua curtipendula, 3 September 1987, A. L.
Hicks, IPL 001774); 2 c? and 1 9 , New Mex-
ico, Socorro Co., Magdalena Mtns., Water
Canyon, 7,000 ft., 16 August 1987, IPL
001756, A. L. Hicks. Deposited BARC, CNC,
KSU, KU, OSU, and USNM.

Remarks. — The aedeagal characters of gi/a
are intermediate between those of doeringae
and pectinata. The styles in gila are less pro-
duced ventrally than in doeringae. The
longer, bladelike styles of gila differentiate
males oi gila from those of pectinata; females
can be readily differentiated on the basis of
the morphology of the female sternum VII
and the bases of the first valvulae. The host of
gila is B. curtipendula. The name gila, a noun
in apposition, denotes the New Mexican
mountains where the holotype was collected.

5. Flexamia bandarita, n. sp.

Length of 6 3.4 (3.3-3.5) mm, 9 3.7
(3.4-4.0) mm; head width of 6 1.09 mm, 9
1.04 mm; head not strongly produced; median
length of crown 0.64 x head width and 1.33 x
interocular width (n = 8 d , 3 9 ).

Color variable, ranging from very dark gray
to pale, with irregular, dark markings on dor-

sum and forewings; face very dark with pale
arcs on upper half; variable in lower half.
Head stramineous in lighter specimens with
paired, transverse lines at midpoint of crown
and obli(}ue markings at rear; venter and legs
with irregular, fuscous markings.

Male. — Pygofer strongly produced ven-
trally in rounded lobe; plates (Fig. 17D)
contiguous on basal half, diverging apically
into narrow V; extending to approximately
3/4 length of pygofer; style (Fig. 17F) heavily
sclerotized, sinuate in ventral aspect, extend-
ing almost to the apex of the aedeagus, gradu-
ally narrowing to acute, ventrally directed,
chelate apex (Fig. 17G) that is closely associ-
ated with sclerotized ridge on inner surface of
each plate; connective in lateral aspect with
dorsal keels narrow; aedeagus (Figs. 17B,C)
symmetrical with slender shaft, straight in
ventral aspect, without apical processes. Apex
flared in caudoventral aspect, but varying
from concave to convex. Dorsal apodemal
process small, not attaining the plane of the
shaft of the aedeagus, appearing suboval in
caudodorsal aspect, with lobelike, converg-
ing, ventral apices. Gonopore minute, sub-
apical on caudoventral surface.

Female. — Sternum VII (Fig. 17H) with in-
ner teeth of the hind margin longer than outer
pair. Ovipositor with basal processes of first
valvulae (Fig. 171) recurved.

Types. — Holotype 6: Brewster County,
Texas, Chisos Mountains, Big Bend National
Park, Texas, 7 August 1987, R. F. Whitcomb
(5,200 ft, Bouteloua curtipendula, IPL
003262). Deposited in USNM. Paratypes:
2 (5 , 1 9 , same collection data; 1 S , Texas,
Marathon Basin, 8 August 1984, deposited
USNM, BARC, KSU, KU; 5 c?, 1 9, Chisos
Mountains, D. J. and J. N. KnuU, 17 July
1946, deposited OSU.

Remarks. — Flexainia bandarita resembles
pectinata, gila, and doeringae, but the sinuate
styles of bandarita separate it from these spe-
cies. The middle teeth of the female sternum
VII are longer than the outer pair, suggesting
an intermediate stage in development of the
median process that is present in doeringae.
Like pectinata and doeringae, bandarita is a
specialist of side-oats grama and, so far as
known, is restricted to that host. Its known
geographic range (Fig. 15) includes the
higher, more mesic grasslands of the Chisos
Mountains in Big Bend National Park and the

240

Great Basin Naturalist Memoirs

No. 12

Fig. 17. Flexamia bandarita, n. sp.; A, aedeagus and connective, lateral aspect; B, aedeagus and connective, dorsal
aspect; C, ape.x of aedeagus, caudoventral aspect; D, male plates and pygofers, ventral aspect; E, male pygofer, lateral
aspect; F, right style, dorsal aspect; G, apex of right style, lateral aspect; H, female sternum VII; I, bases of first valvulae
of female, dorsal aspect.

Marathon Basin north of the park. This region
is rich in endemism presumably derived by
insularization of high grasslands by desertifi-
cation in the surrounding true Chihuahuan
desert. All insects in our recent series, taken
August 1987, are extremely dark; the season
had been one of record rainfall, and vegetative
growth of all plants in the Chisos Mountains
was very lush.

The name bandarita, a noun in apposition,
is one of the common names for side-oats
grama in Mexico.

6. Flexamia doeringae Beamer & Tuthill
Flexamia doeringae Beamer & Tuthill 1934: 3.

Important characters. — Length of 6 3.5
(3.2-3.7) mm, of 9 3.7 (3.3-4.1) mm. Head
produced (median crown length 1.40 x inter-
ocular width; 0.70 X head width) (d n = 20; 9
n = 20). Face ivory with broad, black, basal,
interocular line, occasional dark markings be-

low. [6] Plates (Fig. IIF) broad through most
of their length, then narrowing sharply to
rounded apices. Style with preapical lobe
strongly developed, flattened and produced
posteriorly; aedeagus (Fig. 12F) with protu-
berance near shaft apex close to tip in ventral
aspect. [ 9 ] Sternum VII (Fig. 14F) with inner
teeth strongly produced and closely associ-
ated to form median process; ovipositor with
each first valvula (Fig. 63A) recurved at base;
recurved portion with transverse lobe cepha-
lad of basal extremity.

Geographic distribution. — This species
is known primarily from the mountains (Babo-
quivari, Catalina, Chiricahua, Huachuca, and
Santa Rita ranges) of southeastern Arizona,
but it also occurs in the Mexican highlands of
Sonora and Chihuahua (Fig. 15). There is one
record from Yavapai Co., Arizona (1 6 , Gran-
ite Dell, 10 July 1933, R. H. Beamer).

Biology. — This species is apparently a

1988

Whitcomb, Hicks: Flexamia Revision

241

specialist on side-oats grama, Bouteloua ciir-
tipendula. Because this grass is a dominant in
the mountains of southeastern Arizona and
because many collectors have visited this re-
gion, doeringae is a common component of
large Flexamia collections (Appendix I). The
climate of the southeastern Arizona moun-
tains is similar to that of the Mexican high-
lands but shows a definite Sonoran influence.
Flexamia doeringae may therefore be a Sono-
ran vicariant that separated from the Chi-
huahuan bandarita and the prairie pectinata,
which are also specialists of B. curtipendula.

Oligophagy coefficients. — Gramineae
1.000; Chloridoideae 1.000; Bouteloua 1.000;
Bouteloua curtipendula 0.943 (n = 35).

Remarks and diagnosis. — The range oi do-
eringae does not overlap that of any other
member of the pectinata species group. This
species can be readily distinguished by its
ivory face, the pair of transverse lines on a
pale, well-produced crown, and the structure
of the external genitalia. The female sternum
VII (Fig. 14F) is particularly distinct in having
a medially produced structure consisting of a
pair of long, central lobes and much shorter
lobes on each side. The middle pair of teeth,
in their size and degree of fusion, are maxi-
mally developed in this species.

7. Flexamia collorum, n. sp.

Length of c? 2.9 (2.7-3.0) mm, 9 3.0
(2.8-3.2) mm; head width of 6 0.87 mm, 9
0.89 mm. Crown not strongly produced; me-
dian length of crown 1.53 x head width and
0.68 xinterocular width (c? n = 11; 9 n = 20).

Crown stramineous to light brown, prono-
tum with four faint, wide, brown stripes,
forewings gray with irregular, dark markings.
Face color and markings variable, without in-
terocular line, but usually black in interocular
area and lower area. Usually at least lorae
pale. Crown with pair of oblique markings at
rear, pair of transverse lines at midlength.
Venter usually predominantly dark.

Male. — Pygofer produced ventrally into an
angulate lobe; caudoventral margin sloping
dorsally to a broadly rounded, caudal lobe.
Plates (Fig. 18D) short, extending to 1/2
length of pygofer, fused basally, contiguous
medially for 2/3 length, acute apically. Plates
produced on inner surface at about midlength
into well-developed, sclerotized ridge that
interfaces with style apices. Connective

(Fig. 18A) in lateral view with dorsal keels
narrow, approximately 1/3 height of dorsal
apodeme; apodemal processes as in ban-
darita; style (Fig. 18F) short, abruptly nar-
rowing beyond preapical lobe to avicephali-
form, mesoventrally directed apices (Fig.
18G). Aedeagus (Figs. 18B,C) symmetrical,
shaft slender, without processes, slightly
flared apically in caudoventral aspect.
Gonopore subapical on caudoventral surface.

Female.— Sternum VII (Fig. 18H) with
four teeth on hind margin; outer pair not well
developed, inner pair distinct, but short and
rounded; ovipositor with basal processes of
first valvulae (Fig. 181) recurved.

Types. — Holotype <S: Kimble Co., Texas,
Junction, 14 September 1987, R. F. Whit-
comb and R. C. Chambers (1,300 ft, IPL
003599, Bouteloua uniflora). Deposited
USNM. Paratypes: 16 d, 19 9, same locality;
Sutton Co., Texas, Roosevelt, 14 September
1987, 14 d, 31 9 (1,400 ft, IPL 003593,
003595, 003598). Deposited at KU, OSU,
CNC, USNM, KSU, and BARC.

Remarks. — Flexamia collorum is distinct
from other members of the pectinata species
group, from which it is readily separated by
the short, avicephaliform style apices and by
the extensively developed ridge on the inside
of the male plates. It has been collected at
several localities (Fig. 19) in the Texas hills
(Edwards Plateau), where it appears to spe-
cialize on Nealley grama, Bouteloua uniflora.
Nealley grama differs from side-oats grama in
having single spikelets at the base of the
rachis; it also has a narrow geographic distri-
bution in Texas and Mexico. Although this
species appears to intergrade with curtipen-
dula in the Mexican state of Coahuila (Johnson
1943), it is readily distinguished from it in
central Texas.

The name collorum is an adjective meaning
"of the hills," in recognition of the Texas hills
where it was discovered.

8. Flexamia jacala, n. sp.

Length of 6 3.5 mm; head width of 6 1.08
mm. Crown produced; length of crown 1.42 x
interocular width, 0.75 x head width.

Color gray with irregular, dark markings on
dorsum and forewings; face without black in-
terocular band; crown with oblique, rear and
transverse, midlength markings. Venter and
legs dark with irregular, fuscous markings.

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Fig. 18. Flexamia collorum, n. sp.: A, aedeagus and connective, lateral aspect; B, aedeagus and connective, dorsal
aspect; C, apex of aedeagus, caudoventral aspect; D, male plates and pygofers, ventral aspect; E, male pygofer, lateral
aspect; F, right style, dorsal aspect; G, apex of right style, lateral aspect; H, female sternum VII; I, bases of first valvulae
of female, dorsal aspect.

Male. — Pygofer (Fig. 20D) as in ban-
darita. Plates (Fig. 20G) extend to approxi-
mately 2/3 length of pygofer, fused basally,
contiguous medially for 2/3 of their length,
converging to bluntly rounded apices; male
genitalia similar to those of bandarita, but
styles (Fig. 20E) straight in ventral aspect in-
stead of sinuate, diverging apically; apices
abruptly turning ventrad, appearing avi-
cephaliform in lateral aspect (Fig. 20F),
preapical lobe of style produced laterally and
caudally. Connective (Fig. 20A) in lateral
view with dorsal keels narrow; apodemal pro-
cesses as in bandarita; aedeagal tip (Fig. 20C)
slightly flared in ventral view. Gonopore sub-
apical on caudoventral surface.

Types. — Holotype S: Mexico, Jacala, Hi-
dalgo, 13 August 1936, E. D. Ball. Deposited
inUSNM.

Remarks. — This species is known from a
single male. Nothing is known of its biology. It
is most closely related to bandarita, from
which it can be distinguished by its styles.

which are straight in ventral aspect instead of
sinuate; also, in lateral aspect the apices are
avicephaliform instead of chelate. The name
jacala is a noun in apposition denoting the
type locality.

9. Flexamia niescalero, n. sp.

Length of 6 3.6 (3.5-3.9) mm, 9 3.8
(3.6-4.1) mm; head width of 6 1.07 mm, 9
1.12 mm. Crown moderately produced; me-
dian length of crown 1.42 x head width and
0.66 X interocular width (d n = 6; 9 n = 16).

Color pale with brown markings. Crown
ivory with pair of oblique markings on rear
and pair of transverse lines at midlength;
forewings pale gray with irregular, dark mark-
ings. Face variable, with or (more commonly)
without black interocular band; specimens
without band variably marked on face with
brown or black.

Male. — Pygofer (Fig. 21 E) with posterior
lobe angulate on ventral and caudal margins,

1988

Whitcomb, Hicks: Flexamia Revision

243

Fig. 19. Geographic distribution of mescfl/ero (<
paucijlora and Bouteloua uniflora, respectively.

and collorum (■), with distribution of their hosts, Muhlenbergia

slightly convex dorsally; plates (Fig. 21D) ex-
tending to approximately 4/5 length of
pygofer, fused basally, diverging gradually
along entire length; lateral margins tapering
mesad to acute apices, inner surface with lon-
gitudinal groove and thickened mesal ridge
along 3/4 of length, ending in small, em-
browned tooth. Connective (Fig. 21A) in lat-
eral view with dorsal keels narrow, approxi-
mately 1/4 height of dorsal apodeme;
apodemal processes as in bandarita; styles
(Fig. 21 F) long in ventral aspect, slightly di-
vergent distally, in lateral aspect with preapi-
cal lobe moderately produced and acute. Api-
cal 1/2 of style flattened and minutely serrate
basally, narrowing to ventromesally directed,
calipterate apices (Fig. 21G), much as in ban-
darita. Aedeagus symmetrical, expanded api-
cally in ventral aspect, appearing bluntly sag-
gitate (Figs. 21C, 22), in lateral aspect slightly
indented preapically; gonopore subapical on
caudoventral surface.

Female.— Sternum VII (Fig. 21 H) with
posterior margin notched deeply, middle
teeth very short and deeply inset, median
incision deep, in some specimens divergent at

margin. Ovipositor with bases of first valvulae
(Fig. 211) with sinuate, recurved processes
that converge medially.

Types. — Holotype 6 : Eddy Co. , New Mex-
ico, 23 August 1985. A. L. Hicks and R. F.
Whitcomb, 4,600 ft, IPL 001977, Muhlenber-
gia paticiflora. Deposited in USNM.
Paratypes: 9 d, 16 9, same locality as holo-
type. Deposited at KU, KSU, OSU, CNC,
BARC, and USNM.

Remarks. — Flexamia mescalero is the only
member of the pectinata group that has not
been collected on Bouteloua. The unique
aedeagal apex (Figs. 21C, 22) and female ster-
num VII (Fig. 21 H) are diagnostic for this
species, give it unique status in the group, and
could justify treatment as a distinct group (see
cladogram of Fig. 58). The host of this species
is New Mexican muhly, Muhlenbergia pauci-
flora. Attempts to find other localities for this
species have been unsuccessful.

The name mescalero is a noun in apposi-
tion honoring the Native American Mesca-
lero Apache nation, which inhabited the
mountains of southeastern New Mexico in
the pre-Columbian era and which is today

244

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No. 12

Fig. 20. Flexamia jacala, n. sp.: A, aedeagus and connective, lateral aspect; B, aedeagus and connective, dorsal
aspect; C, apex of aedeagus, caudoventral aspect; D, male pygofer, lateral aspect; E, right style, dorsal aspect; F, apex
of right style, lateral aspect; G, male plates and pygofers, ventral aspect.

establishing high standards for judicious man-
agement of tribal lands.

II. The abbreviata Group

The abbreviata group consists of a single,
widespread species that specializes on
Bouteloua spp. The morphology of the aedea-
gal apex constitutes an autapomorphy that de-
fines the monobasic group.

10. Flexamia abbreviata (Osborn & Ball)

Deltocephalus abbreviatus Osborn & Ball 1897: 206.
Deltocephalus (Flexamia) abbreviatus, DeLong 1926: 33.
Flexamius abbreviatus , DeLong and Sleesman 1929: 83.

Important CHARACTERS. — Length of c? 3.1
(2.8-3.4) mm, of 9 3.2 (2.9-3.6) mm. Head
moderately produced (mean crown length
1.26 X interocular width, 0.61 x head width)
(d n = 20; 9 n = 20). Face without distinct
interocular line (but very rarely with a brown
band contrasting with a pale face). Face often
with parallel, sinuate, brown lines. [6] pygo-

fer (Fig. 62H) strongly constricted, posterior
lobe strongly produced on upper portion of
posterior margin. Plates (Fig. 8E) shield-
shaped, about 0.65 x pygofer length. Aedea-
gus (Fig. 7H) with elongate, anteapical
gonopore on caudoventral surface, apex with
pair of short, truncate processes curved
cephalad, apodemal processes each with
broad, membranous, apical expansion. [9]
Sternum VII (Fig. lOE) with a medial, trun-
cate projection, which has a median incision
that is almost always decorated on either side
by a pair of infuscated spots.

Geographic distribution. — This species
occurs (Fig. 23) from the Canadian prairie
provinces to Montana, Utah, southwestern
Nevada, Wisconsin, Illinois, Oklahoma,
Texas, and Durango, Mexico (1 <5, 27 mi N
LaZarca, 27 October 1981, M. W. Nielson).

Biology. — Because of the dominance of its
Bouteloua hosts, the densities it achieves on
these hosts, and the accessibilitv of its hosts to

1988

Whitcomb, Hicks: Flexamia Revision

245

Fig. 21. Flexainia mescalero, n. sp. : A, aedeagus and connective, lateral aspect; B, aedeagus and connective, dorsal
aspect; C, apex of aedeagus, caudoventral aspect; D, male plates and pygofers, ventral aspect; E, male pygofer, lateral
aspect; F, right style, dorsal aspect; G, apex of right style, caudoventral aspect; H, female sternum VII; I, bases of first
valvulae of female, dorsal aspect.

collectors, this is the most frequently col-
lected Flexamia species (Appendix I). In the
northern and western parts of its range it is a
specialist on blue grama {Bouteloua gracilis),
and the range of this grass limits its distribu-
tion (Fig. 23). In Chihuahuan grasslands and
montane islands of southern Arizona and New
Mexico, it is an inhabitant of mixed Bouteloua
grasslands, where it may occur on blue, hairy
(B. hirsute), or, more rarely, black (B. eri-
opoda) grama. In the eastern prairie where
the grasslands form a mosaic with patches of
eastern deciduous forest, the only known host
is hairy grama; the eastern boundary of the
range of abbreviata is determined by this
host. In the Edwards Plateau of Texas, abbre-
viata occurs on B. hirsuta, B. pectinata, Muh-
lenbergia reverchonii (seep muhly), and per-
haps other chloridoid grasses.

Oligophagy COEFFICIENTS. — Gramineae
1.000; Chloridoideae 0.936; Bouteloua 0.836.
Prairie: Bouteloua 0.780; B. gracilis 0.745

(n = 428); mixed Bouteloua grasslands: B.
gracilis 0.317, B. hirsuta 0.568 (n = 347).

Remarks and diagnosis. — Males and fe-
males of this species can be recognized by the
combination of their small size, brown face
without interocular band, and unique mor-
phologies of the sternum VII or male plates
and pygofer (Fig. 8E). The hind margin of the
female sternum VII (Fig. lOE) contains four
teeth, a condition we believe to be ho-
mologous with similar structures in the pecti-
nata group. Flexamia abbreviata occurs with
curvata (host: Buchloe dactyloides) through-
out much of the range oi Buchloe. Dark male
specimens of curvata are occasionally con-
fused with abbreviata but can be tentatively
recognized without dissection by characters of
the plates and pygofers.

III. The zacate Group

The zacate group consists of two species
that specialize on bush muhly (Muhlenbergia

246

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No. 12

Fig. 22. Scanning electron micrograph of aedeagus oi Flexarnia mescalero: A, lateral aspect (312X); B, ventral aspect
of aedeagal apex (1680X).

porteri). These sister species are distin-
guished on the basis of their posteriorly pro-
duced male pygofers, curved aedeagus lack-
ing apical processes, and elongate, but
rounded male plates. No other Flexarnia spe-
cies has this combination of characters.

Description of the zacate Group

Medium-sized. Length of S 3.2-4.1 mm,
of $ 3.6-4.1 mm. Dorsum brown. Crown
moderately produced. [6] Plates elongate,
but only about 2/3 length of pygofer, fused
basally, narrowing continuously to apices.
Pygofer (Figs. 25E, 621) with posterior lobe
strongly produced posteriorly. Aedeagus and
connective (Fig. 6C) distinctly articulated.
Aedeagus curved in lateral aspect, without
apical processes (Fig. 25A). [ 9 ] Sternum VII

(Figs. 101, L) with deep, median incision.

Key to Species oizacate Group
1. Males 2

— Females 3

2(1). Aedeagus (Fig. 71) narrow in ventral and lateral
aspects; gonopore (Fig. 24) long and narrow,

extending along apical third of shaft

zacate, n. sp.

— Aedeagus (Figs. 6C, 7J) wide, scimitar-shaped
in lateral aspect, diamond-shaped in ventral as-
pect. Gonopore not elongate, near midpoint of
shaft camjonensis Young & Beirne

3(1). Hind margin of sternum Vll (Fig. lOL) with
outer pair of teeth essentially absent, fused with

medial teeth to form median projection

zacate, n. sp.

— Hind margin of sternum VH (Fig. 101) with

outer teeth distinct, rounded

camjonensis Young & Beirne

1988

Whitcomb, Hicks: Flexamia Revision

247

Fig. 23. Geographic distribution oiabbreviata. Western limit is defined by the range oi Boiitelotia gracilis. Eastern
limit is defined by the range oi Botttelotia hirsuta. Host relationships in Mexico are unknown.

1 1 . Flexamia canyonensis Young & Beirne
Flexamia canyonensis Young & Beirne 1958: 18.

Important CHARACTERS. — Length of 6 3.7
(3.5-4.1) mm, of 9 3.9 (3.8-4.1) mm. Head
moderately produced (1.29 x interocular
width; 0.68 X head width) (d n = 4; 9 n = 5).
Face with clypeus and dorsal portion of genae
black. Clypeus, lora, and lower part of genae
pale yellow. [6] Pygofer (Fig. 621) as in ab-
breviata. Plates (Fig. 9C) gradually narrowing
apically to rounded apices. Aedeagus sym-
metrical, scimitar-shaped in lateral aspect, di-
amond-shaped in ventral aspect. Gonopore
on ventral surface of aedeagus, near midpoint
of shaft. [ 9 ] Sternum VII with deep, median
incision on hind margin. Ovipositor with

bases of each first valvula (Fig. 63H) strongly
recurved, processes sinuate in dorsal aspect.

Geographic distribution. — Flexamia can-
yonensis has been collected at Sabino Can-
yon, Arizona (E. D. Ball), and in our own
studies at Kingman, Portal, and Santa Rita
Experimental Range, Arizona (Fig. 26).

Biology. — Our collections of canyonensis
are from bush muhly, Muhlenbergia porteri.
This ^j ass is common in the type locality, and
we assume that it is the sole host. Flexamia
canyonensis appears to be restricted to the
highlands of the Sonoran Desert. This species
and its Chihuahuan sister, zacate, are among
the few Flexamia species to colonize semi-
arid habitats. Perhaps canyonensis is pro-
tected from desiccation by the microclimate

248

Great Basin Naturalist Memoirs

No. 12

Fig. 24. Scanning electron micrograph of aedeagal apex oiFlexamia zacate, ventral aspect (800X).

produced by its host, which grows within pro-
tective shrubs.

Oligophagy coefficients. — Muhlenber-
gia porteri 1.00 (n = 9, 3 series).

Remarks and diagnosis. — Females can be
easily recognized by their distinctive sternum
VII (Fig. 101), which bears a deep, median
incision. Males can be tentatively recognized
by their characteristic face pattern and pro-
duced, flattened pygofer. The aedeagus of
canyonensis is unique.

12. Flexamia zacate, n. sp.

Length of 6 3.5 (3.2-3.9) mm, 9 3.8
(3.6-4.0) mm; head width of 6 1.04 mm, 9
1.08 mm. Crown produced; median length of
crown 1.48 X head width and 0.71 .x interocu-

lar width (d n = 37; 9 n = 12).

Dorsum ivory. Crown with at least trace of
oblique, basal and midlength, transverse
lines. Face without black interocular line,
black in interocular area with at most pale
arcs, bottom of face brown, with irregular,
pale markings. Venter stramineous with ir-
regular markings.

Male. — Pygofer (Fig. 25E) with weakly de-
veloped constriction before posterior lobe,
ventral lobe small, caudoventral margin pro-
duced posteriorly, dorsal margin unevenly
convex. Plates (Fig. 25D) extending to ap-
proximately 2/3 length of pygofer, fused
basally, contiguous medially for 1/2 length,
acute apically; connective in lateral view (Fig.
25A) with dorsal keels narrow, approximately

1988

Whitcomb, Hicks: Flexamia Rkvision

249

bLK

Fig. 25. Flexamia zacate, n. sp.: A, aedeagus and connective, lateral aspect; B, aedeagus and connective, dorsal
aspect; C, apex of aedeagus, caudoventral aspect; D, male plates and pygofers, ventral aspect; E, male pygofer, lateral
aspect; F, right style, dorsal aspect; G, female sternum VII; H, bases of first valvulae of female, dorsal aspect.

1/4 height of dorsal apodeme, with distinc-
tive, minute tooth on ventral surface, anterior
to distinct joint with the aedeagus; apodemal
appendages campanulate in caudal aspect,
with very slender, anteapical processes di-
rected mesad; styles (Fig. 25F) with apex digi-
tate, curving sharply ventrad, each bearing
small preapical tooth on mesal surface; preapi-
cal lobe not produced. Aedeagus symmetri-
cal, shaft slender and cylindrical without pro-
cesses, curving gradually caudodorsad in
lateral aspect, appearing straight in ventral
aspect. Apex slightly flared in caudoventral
aspect, flattened in lateral aspect, with apical
margin concave. Gonopore (Figs. 24, 25C)
long and narrow, extending along apical third
of caudoventral surface of shaft.

Female.— Sternum VII (Fig. 25G) with
posterior margin produced medially, with
deep, median incision; ovipositor with basal

processes of first valvulae (Fig. 25H) with
broad portions close to base.

Types. — Holotype 6: Brewster County,
Texas, Big Bend National Park (Panther Junc-
tion), 26 August 1985 (3,600 ft, R. F. Whit-
comb and A. L. Hicks; IPL 002093). De-
posited USNM. Paratypes: 33 6 , 10 9 , same
locality and date, IPL 002057, 25 August
1985; 13 (?, 4 9, same locahty and date, de-
posited USNM, BARC, CNC, KU, KSU, and
OSU. Thirty-five immature specimens (IPL
002057) have also been deposited with the
paratypes.

Remarks. — This species is a sister to can-
yonensis but can be distinguished from it by
the shape of the aedeagus, which, though
curved, is much narrower in lateral aspect and
is not diamond-shaped in ventral aspect. The
length of the gonopore of zacate also separates
it from canyonensis. Both canyonensis and

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No. 12

Fig. 26. Geographic distribution of canyonensis (•) and zacate (■) and their host, Muhlenbergia porteri.

zacate are apparently specialists on bush
muhly, Muhlenbergia porteri. This grass oc-
curs chiefly in association with thorny shrubs
such SIS Acacia, Prosopis, Larrea, or Opuntia,
which defend it against large herbivores. The
entire volume of the host shrub may be filled
with the tangled culms of bush muhly; even-
tually the host shrub dies, presumably from
competition. Because of its unique habitat,
zacate is difficult to collect without a vacuum-
collecting device. The name zacate is a noun
in apposition denoting various grasses of the
Muhlenbergia or Sporobolus types. Bush
muhly is known in Mexico as "zacate arana '
(spider grass).

IV. The gramniica Group

The grammica group consists of a single,
distinctive species. The mor^Dhology of the
aedeagal apex and the female sternum VII
provides autapomorphies that define the
group.

13. Flexamia grammica (Ball)

Deltocephalus gravimicus Ball 1900; 204.
Deltocephalus (Flexamia) grammicus), DeLong 1926: 37.
Flexamiiis graminicus , DeLong and Caldwell 19.37: 27.

Important CHARACTERS. — Length of 6 4.5

(4.1-5.0) mm, of 9 4.6 (4.3-5.0) mm. Head
not strongly produced (mean crown length
1.08 X interocular width, 0.54 x head width)
(d n = 20; 9 n = 20). Dorsum (Fig. 2C) with
three pairs of longitudinal, dark stripes, me-
dian pair extending from hind portion of the
crown posteriorly through the scutellum and
wings. Face stramineous with single, black
interocular line. [6] Pygofer (Fig. 620) with
narrow, posterior lobe, dorsal more promi-
nent than ventral constriction in lateral as-
pect. Plates (Fig. 8F) broad, contiguous, par-
allel-sided for much of their length, broadly
rounded apically. Aedeagus asymmetrical
with anteapical gonopore on anterodorsal sur-
face, with pair of recurved processes (Fig.
7E), one edentate on the side of shaft near
gonopore, one at the shaft apex. [ 9 ] Sternum
VII (Fig. lOF) broadly and shallowly concave
on posterior margin.

Geographic distribution. — Flexamia gram-
mica occurs in southern Alberta and Sas-
katchewan south through the Great Plains to
Texas (Fig. 27).

Biology. — In the northern part of its
range, grammica is a specialist on prairie
.sandreed, Calamovilfa longifolia. In the
southern part of its range, it specializes on
C. gigantea.

1988

Whitcomb, Hicks; Flexamia Revision

251

Fig. 27. Geographic distribution oi Flexamia grammica, and two hosts, Calamovilfa longifolia (northern prairie) and
C. gigantea (southern prairie and Southwest).

Oligophagy coefficients. — Gramineae
1.000; Chloridoideae 1.000; Calamovilfa spp.
0.872 (n- 47).

Remarks and diagnosis. — The habitus of
grammica is distinctive, as is its size.

V. The curvata Group

The curvata group consists of two closely
related sister species that specialize on buf-
falograss, Buchloe dactyloides. One (curvata)
is adapted to temperate regions, whereas the
other (surcula) is adapted to the subtropical
climate of south Texas and Mexico. In the
following description synapomorphies are
designated [s].

Description of the curvata Group

Small, deltocephaline leafhoppers. Length
of 6 2.4-3.5 mm, of 9 2.6-3.9 mm. Usually
stramineous in color, but occasionally much
darker. [S] Pygofer (Figs. 62K,L) somewhat
produced caudodorsally. Plates (Figs. 9D,E)
short, shield-shaped, about half length of
pygofer. Aedeagus (Figs. 7F,G) [s] with two
pairs of apical processes; style apex [s] foot-
shaped in broadest aspect. [ 9 ] Female ster-

num VII (Figs. 10J,K) with median incision or
notches, with infuscation surrounding or bor-
dering notch or incision.

Key to Species oi curvata Group
1. Males 2

— Females 3

2(1). Gonopore small, subcircular, opening near
aedeagal apex; apical pair of processes short
(Fig. 7G) surcula DeLong & Sleesman

— Gonopore elongate; apical pair of processes
much longer (Fig. 7F) curvata DeLong

3(1). Female sternum VII (Fig. lOK) with completely

infuscated, median incision .... curvata DeLong

— Female sternum Vll (Fig. lOJ) notched, but
without median incision; infuscated on each side

of notches surcula DeLong & Sleesman

14. Elexamia surcula DeLong & Sleesman

Flexamius [sic] surculus DeLong & Sleesman 1929: 99.

Important char.\cters. — Length of S 3.0
(2.6-3.4) mm, of 9 3.4 (2.9-3.7) mm. Head
moderately produced (mean crown length
1.35 X interocular width; 0.68 x head width)
((J n = 20; 9 n = 20). Face stramineous,
except for broad, brown, basal, interocular

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Fig. 28. Geographic distribution oi Flexamia curvata (•) and surcula (A) and their host, Biichloe dactyloides.

line. [6] Pygofer (Fig. 62L) with posterior
lobe strongly produced; style with apical por-
tion foot-shaped, but longer than in curvata.
Plates (Fig. 9E) as in curvata. Aedeagus (Fig.
7G) with pair of recurved, apical processes,
shorter than in curvata, and pair of anteapical
processes. [9] Sternum VII (Fig. lOJ) infus-
cated on either side of shallow notch. Oviposi-
tor with each first valvula (Fig. 631) with basal
portion curved dorsad through more than 90
degrees, recurved portion oblique and cres-
centiform in dorsal aspect.

Geographic distribution. — Subtropical
Texas, and probably coastal prairies of north-
eastern Mexico (Fig. 28).

Biology. — Flexatnia surcula is restricted to
subtropical grasslands. We have collected it
from Buchloe in the Rio Grande Valley in
Cameron and Hidalgo counties of south
Texas. Collection records from December,
January, and February indicate that this spe-
cies does not diapause and probably breeds
more or less continuously throughout the sea-
son as host condition permits.

Remarks and diagnosis. — This species is
closely related to curvata. We have referred a
specimen labeled "paratype" from San Anto-
nio, Texas, in the OSU collection to curvata.

The sternum VII of surcula females lacks the
deep, median incision oi curvata.

15. Flexatnia curvata DeLong

Deltocephalus (Flexamia) ciirvattis DeLong 1926: 34.
Flexamhis ciirvatus, DeLong and Caldwell 1937: 27.

Important characters. — Length of c? 3.1
(2.4-3.5) mm, of 9 3.4 (2.6-3.9) mm. Head
moderately produced (mean crown length
1 . 33 x interocular width; 0. 63 x head width) ( 6
n = 21; 9 n = 23). Face usually pale brown,
often with traces of indistinct, brown interoc-
ular band. [6] Pygofer (Fig. 62K) with poste-
rior lobe simple, somewhat produced cau-
dodorsally. Plates (Fig. 9D) short, shield-
shaped. Style foot-shaped in broadest aspect.
Aedeagus (Fig. 7F) subsymmetrical, recurved
anterior processes extending basad almost to
midlength of shaft, pair of shorter apical pro-
cesses not in bilaterally symmetrical plane;
gonopore an elongate channel on caudoven-
tral surface. Recurved anteapical processes
extending basad almost to middle of shaft;
pair of shorter apical processes extending
basad. [9] Sternum VII (Fig. lOK) with hind
margin concave on each side of median inci-
sion. Ovipositor with base of each first valvula

1988

Whitcomb, Hicks: Flexamia Revision

253

(Fig. 63F) bearing posteriorly directed, slen-
der process.

Geographic distribution. — Flexamia cur-
vata occurs throughout much of the temper-
ate range of its host, Buchloc dactyloides (Fig.
28).

Biology. — This species often occurs in
large populations on Biichloe, where it is a
member of a guild that also includes Athy-
sanella, Pohjamia, Laevicephalus, and Gillet-
tiella species (Whitcomb et al. 1987). In the
subtropical part of the range of bufFalograss
(in south Texas and, presumably, Mexico),
curvata is replaced by surcula.

Oligophagy coefficients. — Gramineae
1.000; Chloridoideae 0.986; Biichloe dacty-
loides 0.925 {n = 560).

Remarks and diagnosis. — This species is
readily recognized by the combination of its
small size and external genitalic characteris-
tics. Most specimens are light stramineous,
with conspicuous, black spots in the middle
and apical cells of the forewing (Fig. 4B); these
specimens are unmistakable. Very large male
specimens of curvata might be confused with
inflata, whose plates and pygofer are superfi-
cially similar. However, inflata, which like
curvata is usually light stramineous in color,
never has a dark spot in the center of the
forewing. Dark specimens of curvata are oc-
casionally referred erroneously to abbreviata.

VI. The picta Group

This group consists of two species, each
with a distinctive habitus, which can be unam-
biguously designated as sister species on the
basis of the morphology of the male aedeagus
and pygofer, and the indistinct articulation
between the aedeagus and connective. In ad-
dition, both species feed on Aristida. One
(pyrops) has, perhaps, the most distinctive
habitus in the genus (Fig. 2E), but picta can
also be recognized without dissection. In the
following description synapomorphies defin-
ing the group are designated [s].

Description of the picta Group

Medium-sized or large, deltocephaline
leafhoppers. Length of 6 2.9-4.0 mm, of 9
2.9-4.9 mm. Grown variably produced (in
pyrops, reaching more than 2 x interocular
width). [ 6 ] Pygofer [s] notched on dorsal mar-
gin, posterior lobe angulate, sharply pro-
duced. Plates small in relation to large pygo-

fers; [s] joint between aedeagus and connec-
tive indistinct. Aedeagus symmetrical, shaft
elongate, [s] apex capitate, pair of slender,
recurved, anteapical processes with entire
edges. Gonopore subapical on caudoventral
surface.

Key to Species of picta Group
1. Head produced, with crown more than twice

interocular width (Fig. 2E) pyrops (Crumb)

— Head only moderately produced . . picta (Osborn)

16. Flexamia picta (Osborn)

Deltocephalus pictus O.sborn 1907: 165.
Deltocephalns funahuhi.s Crumh 1915; 189.
Deltocephalus (Flexamia) pictus, DeLong 1926: .32.
Flexamius [sic] pictus, DeLong and Sleesman 1929: 83.

Important CHARACTERS. — Length of 6 3.5
(2.9-4.0) mm, of 9 3.6 (2.9-4.2) mm. Head
moderately produced (1.38 x interocular
width; 0.64 X head width) (c? n = 24; 9 n =
20). Face pale yellow, with broad interocular
band. Forewings usually dark gray, contrast-
ing with lighter crown and pronotum, dark-
pigmented spot in apex. [ 6 ] Pygofer (Fig. 5A)
similar to that of pyrops. Plates (Fig. 9A) very
short, only about 1/3 length of pygofer, nar-
rowed apically. Aedeagus and connective
with indistinct joint with the connective,
gonopore subapical on caudoventral surface.
Aedeagal apex capitate; expanded portion
with a few small teeth; pair of slender, re-
curved, anteapical processes with entire
edges arising on each side of gonopore. [9]
Sternum VII with median projection. Ovipos-
itor with base of each first valvula (Fig. 63J)
curved through 180 degrees, obliquely bisin-
uate in dorsal aspect.

Geographic distribution. — This species
occurs throughout much of the Southeast,
from Massachusetts to Kansas, and south
through the mixed prairie to Texas (Fig. 29).

Biology. — Flexamia picta appears to be as-
sociated with Aristida spp. Some of the colo-
nized species are annuals. This is a unique
situation in Flexamia hosts; all other known
hosts are perennials.

Oligophagy coefficients. — Gramineae
1.000; Ghloridoideae 0.919; Aristida spp.
0.870 (n = 285).

Remarks and diagnosis. — It is important
for students of the genus to learn the habitus
of picta, which is very common throughout
much of its range. This species can be recog-
nized by the combination of a dark-pigmented

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Great Basin Naturalist Memoirs

No. 12

Fig. 29. Geographic distribution of three Aristida speciahsts; Flexamia arizonensis (O), pyrops (A), and picta (•).

spot in the apex of the forewing (Fig. 4C) in
combination with the face characteristics. The
face is usually sordid yellow but occasionally is
light gray or very pale brown. A few dark
individuals of certain other species (e.g.,
clayi) may have a similar wing spot, but if so,
the face is never pale. The geographical distri-
bution of picta (southeastern) is also usually
indicative. The range of picta overlaps the
ranges of flexulosa and arizonensis, which
have superficially similar faces, only in central
Texas, Oklahoma, and Kansas. Finally, the
small male plates, which contrast with very
large py gofers, are diagnostic for picta. Thus,
dissection is usually unnecessary for recogni-
tion of this species.

17. Flexamia pyrops (Crumb)

Deltocephahis pyrops Crumb 1915: 191.
Deltocephalus (Acurhinus) pyrops, DeLong 1926: 21.
Acurhinus pyrops, DeLong 1948: 226.
Flexamia pyrops, Oman 1949: 167.

Important CHARACTERS. — Length of 6 3.8
(3.6-4.0) mm, of $ 4.2 (3.8-4.9) mm. Head
very strongly produced (crown length 2. 19 x
interocular width, 1.00 x head width) {6 n ==
20; 9 n = 20). Face stramineous, interocular
line with three or more short, dentate projec-

tions from lower margin. [6] Py gofer (Fig.
5B) with distinct notch on dorsal margin, pos-
terior lobe produced ventrad in conspicuous,
acute process; plates (Fig. 9B) very short, no
more than 1/3 length of py gofer, rounded api-
cally. Joint between connective and aedeagus
not well developed. Aedeagus symmetrical,
gonopore on caudoventral surface, pair of re-
curved, anteapical processes arising basad of
gonopore. Apex of shaft (Fig. 7L) capitate,
similar to that of picta, but with fewer, more
distinct teeth. [9] Sternum VII (Fig. lOH)
with hind margin conspicuously concave at
middle.

Geographic distribution. — Although this
is a southeastern species, it occurs as far north
as Pennsvlvania, Ohio, Illinois, and Kansas
(Fig. 29).^

Biology. — Flexamia pyrops, like F. picta,
is closely associated with Aristida spp., but
the two species are seldom found together.
The type oi pyrops from Tennessee was taken
on "Aristida longespica. " In Illinois the host
was A. dichotoma var. curtisii (Whitcomb
1957).

Remarks and diagnosis. — The habitus of
pyrops is unique and diagnostic.

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Whitcomb, Hicks; Flexamia Revision

255

VII. The albida Group

The albida group comprises two species
that are very distinct in general facies and
geographic distribution. Common character-
istics of the species are their patterns of dorsal
stripes, face patterns, and their bifurcate, api-
cal, aedeagal processes (an apomorphy shared
with serrata). The morphology of the male
pygofers of the species may constitute a syn-
apomorphy. The biology of albida and slos-
sonae appears to be more complex than that of
many other Flexamia species. The dorsal
stripes, face patterns, and large size of these
species suggest that they may have arisen
from an ancestor related to Spartopyge.

Description of the albida Group

Medium-sized to large. Length of 6
3.4-4.2 mm, of $ 3.6-4.8 mm. Dorsum with
distinctive pattern of stripes extending from
head across pronotum, scutellum, and
forewings. Face pale with black interocular
band. Male pygofer with poorly differentiated
posterior lobe. Aedeagus with pair of bifur-
cate, apical appendages, unpaired ventral
process.

Key to Species of the albida Group

1. Forewings rounded, male plates (Fig. 8A) not
exceeding pygofer, female sternum VII not
trilobed; prairie albida (Osborn & Ball)

— Forewings obliquely truncate, male plates ex-
ceeding pygofer (Fig. 8B); female sternum VII
trilobed (Fig. lOB); Florida slossonae (Ball)

18. Flexamia albida (Osborn & Ball)

Deltocephalus albidus Osborn & Ball 1897: 201.
Deltocephalus (Flexamia) albidus, DeLong 1926: 36.
Flexamius albidus, DeLong and Sleesman 1929: 83.

Important CHARACTERS. — Length of c? 3.7
(3.4-3.9) mm, of 9 4.3 (3.6-4.8) mm. Crown
not strongly produced (median crown length
1.08 X interocular width, 0.57 x head width)
(6 n = 12; 9 n ^ 21). Face entirely pale.
Pronotum (Fig. 2A) traversed by six longitudi-
nal stripes, central pair extending to rear of
crown, across scutellum, and as spots on base
of forewing; pair of transverse lines midway on
crown. Forewings subhyaline with several
highly pigmented cells, conspicuously pig-
mented, costal cross-veins, and conspicuous
pigmentation along the inner edge of the
forewing. [ 6 ] Pygofer (Fig. 62Q) with poste-
rior lobe short, with small, rounded, ventrally

produced lobe. Plates (Fig. 8A) elongate, un-
notched. Aedeagus symmetrical, gonopore
subapical on caudoventral surface. Aedeagal
apex (Fig. 7B) with pair of distinctive, bifur-
cate processes, each with few teeth on the
dorsal margin. Apodemal processes greatly
expanded at apex. [ 9 ] Sternum VII (Fig. lOA)
with posterior margin shallowly concave on
each side of produced median portion.
Ovipositor with base of each first valvula (Fig.
630) recurved, basal extremity rounded and
directed caudodorsad.

Geographic distribution. — ^This species
occurs in the prairie of Minnesota and North
Dakota, east to Illinois and south to the Black-
land tall-grass prairie of Texas (Fig. 30).

Biology. — Flexamia albida is a resident of
tall-grass and mixed prairies. Many records
are from Schizachyrium scoparium. How-
ever, albida is not found in pure stands of S.
scoparium, suggesting a life history more
complex than that of other Flexamia species.

Oligophagy coefficients. — Gramineae
1.000; Panicoideae 0.674 (Schizachyrium sco-
parium 0.674); Chloridoideae 0.196 (Boute-
loua curtipendula 0. 196) (n = 46).

Remarks and diagnosis. — The habitus of
F. albida is unique, and its geographic range
does not overlap that of the other striped Flex-
amia species.

19. Flexamia slossonae (Ball)

Deltocephalus slossoni [sic] Ball 1905: 119.
Deltocephalus (Flexamia) slossoni, DeLong 1926: 36.
Deltocephalus (Secopennis) slossoni, DeLong and Slees-
man 1929: 85.
Flexamia slossoni, DeLong and Caldwell 1937: 27.
Secopennis slossonae, Oman 1949: 168.

Important characters. — Length of d 3.8
(3.5-4.2) mm, of 9 4.2 (3.8-4.5) mm. Crown
produced (median crown length 1.38 x inter-
ocular width, 0.64 X head width) (c? n = 20; 9
n = 20). Face ivory with black interocular
band at base. Dorsum with longitudinal lines
extending from rear of the head across prono-
tum and* scutellum to rear of forewings (Fig.
2B). Male plates (Fig. 8B) exceeding pygofer
in length. [6] Aedeagus (Fig. 7C) symmetri-
cal, gonopore an elongate slit extending
through most of length of unpaired ventral
process; apical processes bifurcate, without
teeth. [ 9 ] Sternum VII (Fig. lOB) conspicu-
ously tri-lobed.

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Fig. 30. Geographic distribution oiFlcxamia serrata (■), alhida (•), slossonae (D), and ritana (O).

Geographic distribution. — Flexamia slos-
sonae is restricted (Fig. 30) to subtropical
grasslands of south Florida.

Biology. — DeLong (1926) reported find-
ing adults and nymphs of slossonae on salt-
grass (Distichlis spicata). We have found the
species on the Muhlenhergia prairies of the
Florida Everglades, in the absence of Distich-
lis. It seems likely that this species is a hal:)itat
rather than a host specialist.

Remarks and diagnosis. — Even if slos-
sonae were not confined to the subtropical
region of .south Florida, its habitus would per-
mit its unambiguous identification. The dorsal
stripes of alhida (Fig. 2A) are somewhat simi-
lar to those of slossonae, but in the latter
species the stripes extend conspicuously to
the rear of the forewings (Fig. 2B). Also, the

forewings (Fig. 2B) are obliquely truncate,
exposing the terminal abdominal segment,
which is also striped. The forewings of ritana
(Fig. 2D) are broadly truncate.

VIII. The ritana Group

The ritana group consists of a single spe-
cies. The monobasic group is defined by the
bizarre aedeagal apex, which we designate as
an autapomorphy.

20. Flexamia ritana Beamer

Flcxainiii ritana Beamer 1936: 257.

Important CHARACTERS. — Length of 6 4.2
(3.9-4.4) mm, of 9 4.4 mm. Crown produced
(mean crown length 1.67 x interocular width;
0.73 X head width) (d n = 4; $ n = 1). Dorsum

1988

Whitcomb, Hicks; Flexamia Revision

257

with stripes extending from head to forcwings
(Fig. 2D). Face pale white with broad, hiscous
interocular band. [6] Plates (Fig. 8D) about
1/2 length of pygofer, fused basally, tapering
to bluntly acute apices. Aedeagus symmetri-
cal, gonopore on caudoventral surface at base
of unpaired processes (Fig. 7D); processes
with entire edges, together lyriform; apode-
mal processes with anteapical protuberances.
[ 9 ] Sternum VII with conspicuous, median
projection on hind margin.

Geographic distribution. — Flexamia ri-
tana has been collected only in the Santa Rita
Mountains of Arizona (Fig. 30).

Biology. — Despite many field hours ex-
pended in search, ritana has not been found
since its collection in 1935. The original field
notes of R. H. Beamer are not explicit in
pinpointing the locality (see notes, Appendix
III). Our field studies have emphasized col-
lections on relatively pure patches of grass.
Considering the accumulating evidence that
primitive Flexamia species may be habitat
specialists, future efforts should perhaps be
directed at diverse habitats rather than single
host species.

Remarks and diagnosis. — Superficially,
ritana (Fig. 2D) resembles albida (Fig. 2A)
but lacks transverse lines on the crown and
has no darkly pigmented forewing cells. The
forewings (Fig. 2D) are broadly truncate. The
male plates and female sternum VII are en-
tirely unlike those of either slossonae or al-
bida.

IX. The serrata Group

This group consists of a single species. The
defining autapomorphy of the group is the
pair of dorsal, aedeagal processes.

21. Flexamia serrata Beamer & Tuthill
Flexamia serrata Beamer & Tuthill 1934: 4.

Important characters. — Length of 6 3.9
(3.6-4.3) mm, of 9 3.8 (3.2-4.4) mm. Crown
moderately produced (mean crown length
1.26 X interocular width, 0.66 x head width)
{6 n = 20; 9 n = 20). Face pale with interocu-
lar band consisting of parallel lines inter-
rupted at middle. [6] Pygofer (Fig. 62P) with
posterior margin of lobe subangulate above
midlength. Plates (Fig. 8C) short, broadly tri-
angulate, about half length of pygofer, acute at
apices. Aedeagus (Fig. 7A) subsymmetrical.

gonopore anteapical on caudoventral surface.
Prominent, unpaired, ventral process basad
of gonopore, extending basad along shaft, pair
of shorter, recurved processes arising on each
side of gonopore. Apodemal processes with
mesal, anteapical protuberances. The un-
paired aedeagal processes of some males from
Linton, North Dakota (11 August 1977, 3 6,
3 9, R. F. Whitcomb, IPL 000351, and 24
July 1985, 34 c? , 16 9 , R. F. Whitcomb and
E. A. Clark, IPL 001612) and Balta, North
Dakota (1 6, 3 August 1985, K. G. A. Hamil-
ton) are broken; such genitalia can be initially
confusing. [9] Sternum VII (Fig. IOC) with
broad, median, posterior projection. Oviposi-
tor with base of each first valvula very strongly
recurved, recurved portion approximately
parallel to axis of valvula (Fig. 63 P).

Geographic distribution. — This species
(Fig. 30) occurs from the southern tier of west-
ern Canadian provinces to Utah and North
Dakota.

Biology. — Flexamia serrata is apparently
associated with Muhlenbergia richardsonis ,
where it often occurs (especially in Canada)
with decora.

Oligophagy coefficients. — Gramineae
1.000; Chloridoideae 1.000; Muhlenbergia
richardsonis 1.000 (n = 57).

Remarks and diagnosis. — The face pattern
of serrata (Fig. 3A) is unique. The range of
this species overlaps considerably with that of
stylata, which has superficially similar facial
lines. However, in stylata the lines are not
interrupted at the middle and contrast with a
yellow rather than white face. The male plates
oi stylata (Fig. 34E) and the unique morphol-
ogy of the male pygofer (Fig. 5C) also differen-
tiate it from serrata. Whereas all individuals
of stylata have at least a trace of a transverse
line midlength on the crown, serrata has a
completely unmarked crown.

X. The flexulosa Group

The flexulosa species group consists of
11 species. Evolution within this group has
occurred on Aristida, Bouteloua, and, espe-
cially, Muhlenbergia. We have found no syn-
apomorphy to define the group, but we have
been able to define four subgroups. The nomi-
nate subgroup consists of two species. One of
these, arizonensis, specializes on Aristida
purpurea and/or other Aristida species. The
other member of this subgroup is flexulosa, a

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No. 12

blue grama (Bouteloua gracilis) specialist.
This species is designated as a sister to ari-
zonensis on the basis of the extremely short,
apically divergent plates, which are synapo-
morphic. The decora subgroup consists of
three species. F. decora and youngi are iden-
tified as sisters on the basis of their ventral,
aedeagal processes, which are much longer
than the paired processes. Both are specialists
on Muhlenbergia richardsonis . The current
geographic distribution of these species sug-
gests that they represent vicariant segregates
of an ancestral Muhlenbergia specialist. The
third species, modica, is a specialist on Muh-
lenbergia repens (a close relative of M.
richardsonis ) in more mesic grasslands of the
eastern Desert Plains. This species is linked to
the decora subgroup by the morphology of the
male pygofer and female sternum VII. The
third subgroup consists of three species:
arenicola, a specialist on Muhlenbergia pun-
gens; stylata, a specialist on Muhlenbergia
species in uplands of the northern prairie and
the eastern Rocky Mountains; and celata, a
specialist of chloridoid grasses in sandhills,
including sand blowout grass, Redfieldia flex-
uosa. The latter two species are designated
sisters on the basis of their unique male
pygofers, which are synapomorphic. A fourth
subgroup consists of three species. One of
these, inflata, a species of moist northern
prairies, commonly occurs on Muhlenbergia
asperifolia, which enables it to colonize ripar-
ian areas of the Southwest. Flexamia texana, a
rare species from south central Texas whose
biology is unknown, is designated sister to
beameri, a species from New York whose biol-
ogy is also unknown. The triangulate male
plates and boat-shaped pygofers are synapo-
morphic for this subgroup.

Description oi the flexulosa Group

Small to large. Length of 6 2.9-4.6 mm,
of 9 3.0-4.7 mm. Gray to stramineous.
Crown not strongly produced in some species
{decora, inflata), moderately produced in
others. Face in many species iflexulosa and
decora subgroups) white, ivory, or at the
darkest, very pale yellow, contrasting sharply
with dark, usually black interocular band.
Face patterns in stylata subgroup similar, but
with interocular band of parallel, discrete
lines. In other species {inflata subgroup), face
stramineous with no interocular band or at

most indistinct band not contrasting with
lower face. Male plates triangulate, short or
elongate, often apically divergent, in some
species fused basally. Aedeagus and connec-
tive articulated. Aedeagus with paired apical
processes and ventral unpaired process in
sagittal plane, or displaced from sagittal plane
{inflata subgroup). Gonopore apical or sub-
apical on caudoventral surface. Female ster-
num VII with median projection.

Key to Males of the/7erH/osa Group

1. Pvgofer produced caudallv in spinelike process

(Fig. 5C) ■ 2

— Pygofer not so 3

2(1). Shaft of aedeagus thick, conspicuously curved,
expanded apicallv with long, apical appendages
(Fig. 31H) stijlata (Ball)

— Shaft of aedeagus slender, not conspicuously
curved or expanded apically, with shorter apical
appendages (Fig. 311) . . . celata Lowry & Blocker

3(1). Pygofer with posterior lobe produced, heavily
sclerotized, and angled on ventral margin (Figs.
32E; 62MM) arizonensis Young & Beime

— Pygofer with posterior lobe rounded ventrally . . 4

4(3). Aedeagus with greatly elongate, unpaired ven-
tral process and with shaft straight or arched
caudodorsally 5

— Aedeagus with paired and unpaired processes
more or less same length or if one greatly elon-
gate, then shaft sigmoidal 6

5(4). Aedeagal shaft (Fig. 31G) curved, unpaired pro-
cess straight decora Beamer & Tuthill

— Aedeagal shaft (Figs. 31F, 38A,B) straight, un-
paired process recurved youngi, n. sp.

6(4). Aedeagal shaft at least weakly sigmoidal in lat-
eral aspect 7

— Aedeagal shaft arched caudodorsally, not sig-
moidal 9

7(6). Aedeagus with paired processes less than 1/4
length of unpaired process, and not diverging
from plane of shaft in lateral aspect (Fig. 31E)
inflata (Osborn & Ball)

— Paired processes longer, and diverging from
plane of shaft in lateral aspect 8

8(7). Aedeagal processes irregular, apex strongly
asymmetrical (Fig. 31A); south central Texas
texana Young & Beirne

— Aedeagal apex subsymmetrical (Fig. 31 D), pro-
cesses regular; New York beaineri, n. sp.

9(6). Pygofer in lateral aspect with caudoventral mar-
gin from ventral lobe to caudal apex straight, not
curved; paired processes of the aedeagus in lat-
eral aspect diverging dorsally from the plane of
the shaft; dorsal keels of the connective narrow;
plates (Fig. 33A) small, short flextdosa (Ball)

1988

WhITCOMB, HlCKS: FLEXAMIA REVISION

259

Fig 31. Aedeagal apices of the Flexamia flexulosa group: A, texana- B, imputans; C, areolata; D,beamen; E, inflata;
F youngU G, decora; H, stylata; I, celata: J, arenicola; K, i?exu/o5a, L, arizonensis; M, modtca. Aspects: CL,
caudolateral CV, caudoventral; DS, dorsal; LT, lateral; VN, ventral. Redrawn from Young and Beime (1958).

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DlX

Fig. 32. Male plates and pygofers oi some flcxulosa group species, lateral aspect: A, ijoungi, B, mocUca, C, arenicola,
D, decora, E, arizonensis, F,fJexuIosa.

— Pygofer in lateral aspect with caudoventral margin
curved from ventral lobe to caudal apex; paired
processes of aedeagus in lateral aspect diverging
weakly or not at all from the plane of the shaft;
dorsal keels wider in lateral aspect; plates (Figs.
33C,F) comparatively long 10

10(9). Apices of paired processes curving ventrad in
lateral aspect; processes and shaft of aedeagus
comparatively robust . . . inodica Beamer & Tuthill

— Apices of paired processes curving dorsad in
lateral aspect; processes and shaft comparatively
slender arenicola Lowry & Blocker

Key to Females oi the flexulosu Group
1. Ovipositor with base of each first valvula curved

2(1).

dorsad and laterad, forming angle of more than
45 degrees with long axis of valvula and extend-
ing laterad beyond lateral margin of valvula
when viewed from above (Figs. 63Q,R,AA) .... 2

Ovipositor with recurved portion of each first
valvula not extending laterad beyond lateral
margin of valvula (or only very slightly so); if
longer than broad, then with long axis forming
angle of less than 45 degrees with axis of valvula
in dorsal aspect 4

Recurved base of each first valvula very slender
in dorsal aspect, posterior margin concave (Fig.
63AA) stylata (Ball)

Recurved base of each first valvula broader in

1988

Whitcomb, Hicks: Flexamia Revision

261

Fig. 33. Male plates and pygofers of the Flexamia flexulosa group: A, flexulosa, B, arizonensis; C, arenicola; D,
decora; E, youngi; F, modica.

dorsal aspect, anterior margin straight or convex 3

3(2). Recurved base of each first valvula more distant
from broadest part of valvula and with dorsal
portion often lightly sclerotized and with small,

anterior point (Fig. 63Q)

decora Beamer & Tuthill

— Recurved base of each first valvula without
small, anterior point and closer to broadest part

of valvula (Fig. 63R) . . . modica Beamer & Tuthill

4(1). Recurved portion of each first valvula short,
with acute, dorsal projection directed anterolat-
erad (Fig. 63T) inflata (Osborn & Ball)

— Recurved portion of first valvula without such
projection 5

5(4). Recurved portion of first valvula concave poste-
riorly in dorsal aspect (Fig. 63Y)

arizonensis Young & Beirne

— Recurved portion of each first valvula not so ... 6

6(5). Recurved portion of each first valvula directed

almost caudally (Fig. 63BB) flexulosa (Ball)

— First valvula not so 7

7(6). Upper New York State beameri, n. sp.

— South central Texas texana Young & Beirne

22. Flexamia arizonensis Young & Beirne

Flexamia arizonensis Young & Beirne 1958: 30.

Important CHARACTERS. — Length of 6 3.3
(3.0-3.8) mm, of 9 3.6 (3.2-4.0) mm. Head
variably produced (1.40 x interocular width;
0. 70 X head width) {6 n = 20; 9 n = 20). Face
pale white with black interocular band. [6]
Pygofer with posterior lobe (Fig. 62MM) api-
cally truncate, ventral portion extending ven-
trad as short, heavily sclerotized process.
Plates short, less than 1/2 length of pygofer,
apically divergent. Aedeagus symmetrical,
paired apical processes arising on each side of
subapical gonopore, unpaired ventral process
about one-third length of shaft. [ $ ] Sternum
VII with broad, median projection. Oviposi-
tor with each first valvula (Fig. 63Y) curved at
base more than 90 degrees, recurved portion
concave caudally in dorsal aspect.

Geographic distribution. — ^This species
is more widely distributed than had been pre-
viously supposed, occurring (Fig. 29) from

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Fig. 34. Male plates and pygofers of the Flexamiaflexulosa, imputans, and areolata groups, ventral aspect: A, inflata;
B, texana; C, heameri, D, celata; E, stylata; F, imputans; C, areolata.

western Nebraska, Colorado, Utah, central
Texas, and Arizona to northern Mexico
(Sonora and Chihuahua). Two females from
southern California (Pine Flats Camp, Indio,
L. H. Banker, 12 July 1941 [KU]) may repre-
sent this species.

Biology. — This species is a specialist on
perennial Aristida spp., especially A. pur-
purea.

Olicophagy coefficients. — Gramineae
1.000; Chloridoideae 1.000; Aristida spp.
0.932.

1988

Whitcomb, Hicks: Flexamia Revision

263

{

\ -"1""'

iL'^'--'

—?f.-

'^'":"""r"'

■ 7 .

J— -'-'''

Fig. 35. Geographic distribution o( Flexamia flexulosa and its host, Bouteloua gracilis.

Remarks and diagnosis. — Prior to the
study of Young and Beirne, arizonensis had
been confused with flexulosa. Males of these
species can be distinguished by the lateral
aspect of their undissected pygofers (Fig.
32E,F; see discussion under flexulosa). Al-
though the dorsum of arizonensis is usually
pale, several specimens from southern Ari-
zona show dorsal stripes that are reminiscent
of those ofritana. Since the external genitalic
characteristics of arizonensis are very distinct
from those oi ritana, such specimens pose no
problem for identification. However, they are
important in suggesting an evolutionary link
between the ritana and flexulosa groups.

23. Flexamia flexulosa (Ball)

Deltocephalus flexulosus Ball 1899: 189.
Deltocephalus (Flexamia) flexiilosus , DeLong 1926; 31.
Flexamius flexulosus, DeLong and Sleesman 1929: 84.

Important CHARACTERS. — Length of S 3.2
(2.9-3.7) mm, of 9 3.5 (3.0-4.1). Head vari-
ably produced (1.26 x interocular width; 0.60
xhead width) (d n = 40; 9 n = 44). Face pale
with black interocular stripe. [6] Pygofer
(Fig. 62NN) with posterior lobe rounded api-
cally. Plates very short, less than 1/2 length of

pygofer, apically divergent. Aedeagus sym-
metrical, gonopore anteapical on caudoven-
tral surface; paired apical processes short,
minutely serrulate along one edge; unpaired
ventral process short, grooved, less than one-
fourth length of the shaft, acute at apex. [ 9 ]
Sternum VII with hind margin produced to
form convex lobe; ovipositor with each first
valvula curved through more than 90 degrees
at base, recurved portion slender, almost par-
allel to long axis of valvula, with narrow pro-
jection extending caudad at basal extremity.

Geographic distribution. — This species
occurs (Fig. 35) from eastern Nevada, Utah,
Wyoming, Montana, and the southern tier of
Canadian provinces from Alberta to Mani-
toba, south through the Great Plains to west-
ern Texas, Arizona, and northern Mexico
(Chihuahua).

Biology. — Although flexulosa and ahhre-
viata are both specialists of blue grama
(Bouteloua gracilis), their ranges do not com-
pletely overlap. Whereas y/(?x«/osa appears to
be adapted solely to blue grama in cool, dry
climates, abbreviata is adapted to warmer
grasslands, especially the mixed Bouteloua

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Great Basin Naturalist Memoirs

No. 12

grasslands of Texas, New Mexico, and Ari-
zona. Such differences may be important
clues to the evolutionary origins of these Flex-
amia species.

Oligophagy coefficients. — Gramineae
1.000, Chloridoideae 0.973; Bouteloua gra-
cilis 0.889 (n - 380).

Remarks and diagnosis. — Young and
Beirne (1958) consider flexulosa to be "greatly
variable." Actually, we have found substantial
variability in all Flexamia species for which
large sample sizes were available; flexulosa
seems to us to be no more or less variable than
other species. The face pattern of flexulosa
(black interocular band contrasting with a pale
face) is, in fact, remarkably stable. Unfortu-
nately, this contrasting pattern is shared by
other members of the flexulosa group (ari-
zonensis, modica, arenicola, decora, and
youngi). Although the definitive identity of
these species should be ascertained by exami-
nation of genitalic characteristics, flexulosa is
an abundant species, so it is important to de-
fine external characters that permit it to be
sorted from less common species. The lateral
aspect of the male pygofer (Figs. 32A-F) is a
useful character in sorting white-faced mem-
bers of the flexulo.sa group. In flexulosa the
posterior margin of uncleared male pygofers
appears rounded. The pygofer (Fig. 32E) of
the widely distributed arizonensis is truncate
apically, but it is necessary to clear the speci-
men to see the distinctive, heavily sclerotized
ventral process. In regions where both decora
SLud flexidosa occur, decora can often be rec-
ognized by its characteristic habitus (see dis-
cussion under that species), and by its pygofer
(Fig. 32D), which appears narrower than that
of other white-faced species. The pygofer of
rnodica (Fig. 32B), an uncommon but locally
abundant (New Mexico mountains) specialist
of Muhlenbergia repens, is also diagnostic.
The pygofer of arenicola (Fig. 32C) is also
more or less diagnostic; this insect is an inhab-
itant of Muhlenbergia pungens in the Ne-
braska Sand Hills and the Four Corners re-
gion of the Colorado Plateau (Fig. 36). Thus,
the combination of male pygofer, geographic
range, and host data (if available) provides an
excellent character set that permits tentative
recognition of pale-faced males of the flexu-
losa group. Nevertheless, the identity of
these species, induding flexulosa , should be

confirmed by examination of characters of the
genitalia.

24. Flexamia arenicola Lowry & Blocker
Flexamia arenicola Lowry & Blocker 1987; .59.

Important characters. — Length of d 3.7
(3.5-3.9) mm (Sand Hills form), 3.4 (3.2-3.8)
mm (Anasazi form); of 9 4.0 (3.8-4.3) mm
(Sand Hills form), 3.7 (3.3-4.1) mm (Anasazi
form). Crown moderately produced (median
crown length 1. 19 x [Sand Hills form] or 1.35 x
[Anasazi form] interocular width; 0.63 x head
width [both forms]) ((5 n - 20, 9 n - 7 [Sand
Hills form]; d n = 20, 9 n = 18 [Anasazi
form]). Crown moderately produced (median
crown length 1.25 x interocular width; 0.67 x
head width). Face pale, with conspicuous,
black interocular band. [6] Pygofer (Fig.
62LL) broadly ovate; plates (Fig. 33C) broadly
triangulate, diverging apically. Aedeagus
(Fig. 31J) symmetrical with gonopore anteapi-
cal on the caudoventral surface, lateral, apical
processes approximately one-fourth shaft
length and curved laterodorsad; unpaired
process approximately the same length and
directed anteriorly. [ 9 ] Sternum VII with
posterior margin produced medially with
small, median notch. Ovipositor with re-
curved process at base of first valvula (Fig.
63S) not exceeding lateral margin, process ex-
tending caudad, sinuate and digitate apically.

Geographic distribution. — There are two
disjunct populations of arenicola (Fig. 36).
One population occurs in the sandhills of
western Nebraska and northeastern Colo-
rado. The other population occurs in the Colo-
rado Plateau and adjacent regions of New
Mexico. The disjunct distribution of arenicola
can be readily explained by the essentially
disjunct distribution of its host (Fig. 36).

Biology. — Flexamia arenicola has been
found almost exclusively on sandhill muhly,
Muhlenbergia pungens.

Oligophagy coefficients. — Gramineae
1.000; Chloridoideae 1.000; Muhlenbergia
pungens 0.989 {n 91).

Remarks and diagnosis. — The unpaired
processes of the aedeagus in males of the Four
Corners population (which we term the
"Anasazi form" in honor of the pre-Columbian
residents of the area) is uniformly absent, but
on close inspection it can be seen to be bro-
ken; this presumably occurs during copula-
tion. The occurrence of the Anasazi form is

1988

Whitcomb, Hicks: Flexamia Revision

265

Fig. 36. Geographic distribution of Flexamia arenicola (Sand Hills
Muhlenbergia pungens.

[•] and "Anasazi" [■] forms) and the host.

patchy, and populations usually are low. Be-
cause the region occupied is now undergoing
desertification and salinization as a result of
climatic changes during the interglacial era,
this form may be a relict population that, like
the host population, is currently stressed by
climatic change. Presumably, the outcome of
these selective pressures could lead to extir-
pation or to evolution of a new species that
would be better adapted to semiarid environ-
ments. The characteristics of the male pygofer
(Fig. 32C) in lateral aspect, in combination
with the face characteristics, are diagnostic for
arenicola. Lowry and Blocker (1986) distin-
guished arenicola from flexiilosa by the ex-
panded dorsal keels of the arenicola connec-
tive, the more robust aedeagal shaft, and the
longer aedeagal processes. They distin-
guished arenicola from modica by its larger
size and nearly symmetrical aedeagus with
lateral, dorsally curved, apical processes. In
addition, the pygofer of modica (Fig. 32B) is
very different from that of arenicola. Some
characters in arenicola are similar to those of
celata and stijlata, but the general facies of the
latter two species separate them from areni-
cola.

25. Flexamia celata Lowry & Blocker

Flexamia celata Lowry & Blocker 1987; 57.

Important CHARACTERS. — Length of d 4.1
(3.9_4.4) mm, of 9 4.5 (4.4-4.7) mm. Crown
not produced (1. 13 x interocular width; 0.58 x
head width) (d n = 19; 9 n = 6). Face, as in
stijlata, pale with broad interocular band con-
sisting of parallel, discrete lines. [6] Pygofer
(Fig. 62PP) with central margin strongly pro-
duced to form spinelike process, as in stijlata.
Plates (Fig. 34D) extending to approximately
2/3 length of pygofer, fused mesally for 1/2
length, narrowed apically to rounded, lateral
lobe, apices meeting in V-shaped notch.
Aedeagus (Fig. 311) symmetrical, shaft slen-
der, neither conspicuously curved nor ex-
panded apically, ventral and paired apical
processes approximately equal in length and
less than 1/4 length of shaft; gonopore apical
on caudoventral surface. [ 9 ] Sternum VII
with posterior margin medially produced,
with slight median notch; ovipositor with
basal processes of first valvulae recurved and
extending laterad beyond lateral margin.

Geographical distribution. — Nebraska
Sand Hills and southwestern Kansas (Fig. 37).

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No. 12

Fig. 37. Geographic distribution ofFlexamia stylata (•) and celata (■) and a host oicelata, Redfleldia flexuosa.

Biology. — This species occurs on sand
blowout grass, Redfieldia flexuosa, the pri-
mary colonist of bare sand in the Sand Hills. It
may also occur on other sandhill grasses. It has
been recorded from Calmyiovilfa longifolia in
southwestern Kansas (I (?, 3 9 , Garden City,
Finney County, 12 June 1948, R. H. Beamer).

Oligophagy coefficients. — Gramineae
1.000; Redfieldia flexuosa 0.612; Muhlenber-
gia pungens 0.265 (n = 49).

Diagnosis and remarks. — This is a close
sister species with stylata, under which diag-
nostic characters have been discussed. The
present-day area of endemism was apparently
created by the southern interface of the ice
sheet during the last glacial maximum (Wright
1970), suggesting that celata could represent a
recent speciation. On the other hand, its mor-
phological characters are much closer to the
ground plan for the flexulosa group than are
those oi stylata. The aedeagal symmetry and
wide pygofer of celata suggest a relationship
with arenicola, which occurs with it in the
Nebraska Sand Hills.

26. Flexamia stylata (Ball)
Deltocephalus stylatus Ball 1899: 190.

Deltocephaltis (Flexamia) stylatus, DeLong 1926: 34.
Flexamius stylatus, DeLong and Sleesman 1929: 84.

Important CHARACTERS. — Length of c? 4:1
(3.7-4.6) mm, of 9 4.2 (3.8-4.6) mm. Crown
not strongly produced (1.17 x interocular
width; 0.54 X head width) (c? n = 33; 9 n =
33). Face pale, except for broad interocular
band consisting of discrete, parallel lines. [6]
Plates (Fig. 34E) with conspicuous, sub-
rectangular apices; pygofer (Fig. 6200) with
posterior margin strongly produced, gradu-
ally narrowed, forming curved, spinelike pro-
cess. Aedeagus asymmetrical; shaft subcylin-
drical; gonopore subapical, at base of three
processes on caudoventral surface; pair of api-
cal processes directed laterobasad, unpaired
ventral process extending basad to midlength
of shaft, but not parallel to axis of shaft, the
three processes appearing subequal in length
in caudoventral aspect; paired processes ap-
pearing shorter in lateral aspect, each slightly
serrate on dorsal edge. [ 6 ] Sternum VII as for
group; ovipositor with base of each first
valvula (Fig. 63AA) curved dorsad through
slightly more than 90 degrees, recurved por-
tion extending laterad beyond lateral margin

1988

Whitcomb, Hicks; Flexamia Revision

267

of valvula in dorsal aspect.

Geographic distribution. — Southern tier
of Canadian provinces south through the
western Great Plains, eastern Rocky Moun-
tains, northern New Mexico, and Arizona
(Fig. 37).

Biology. — This species has been collected
from Muhlenhergia on upland slopes in Mon-
tana and Wyoming. In northern New Mexico
the host is Muhlenhergia wrightii. This host
may also account for records from northern
Arizona.

Diagnosis and remarks. — The male plates
of this species are unique. The face pattern
(interocular bands with discrete lines) that
stylata shares with celata is very distinctive;
superficially similar species almost always
have bands with coalesced lines. Finally, sty-
lata specimens are larger than those of most
other species. Females of stylata and celata
are similar but can be distinguished by charac-
teristics of the bases of the first valvula of the
ovipositor. Also, stylata is a widely dis-
tributed Muhlenhergia specialist, whereas
celata is restricted largely to the Nebraska
Sand Hills, where it is a rare inhabitant of
Redfieldia flexiiosa and other chloridoid
grasses.

27. Flexainia decora Reamer & Tuthill

Flexamia decora Beamer & Tuthill 1934: 2.

Important characters. — Length of c? 3.6
(3.0-4.2) mm, of 9 3.9 (3.4-4.2) mm. Head
not strongly produced (median crown length
1.07 X interocular width, 0.62 x head width)
(c? n = 20; $ n = 20). Face pale with broad
black interocular stripe. [6] Pygofer (Fig.
62FF) with slightly truncate rear margin.
Plates (Fig. 33D) broadly divergent, with nar-
row apices. Aedeagus (Fig. 31G) symmetrical.
Gonopore anteapical on caudoventral surface;
pair of retrorse processes arising on each side
of gonopore, each serrate on dorsal edges.
Unpaired ventral processes, with entire
edges, extending basad more than half length
of shaft. [ 9 ] Sternum VII with hind margin
concave at each side of median convex projec-
tion, notched apically, with teeth on each side
of notch. Ovipositor with each first valvula
(Fig. 63Q) with base expanded dorsally and
laterally.

Geographic distribution. — Flexamia de-
cora, as defined herein, is a specialist oi Muh-

lenhergia richardsonis in the New Mexico
and Colorado Rockies north to the Peace
River region of northwestern Alberta, and
east to Manitoba and North Dakota (Fig. 39).
Its host occurs in eastern North America but
does not occur south of Maine. We have ex-
amined specimens reported by Young and
Beirne (1958) from Kentucky and Virginia.
The Kentucky specimen is a female referable
to inflata. The Virginia specimen labeled only
"Oct. 12, Va., ED. Ball" appears to have been
recently relabeled (and therefore possibly
mislabeled), perhaps at the time of acquisition
of the Ball collection. There are therefore no
reliable records for decora east of the mixed-
grass prairie.

Biology. — Flexamia decora appears to be
largely a specialist of Muhlenhergia richard-
sonis. Large populations are not uncommon,
and decora is the most abundant Flexamia
species in the CNC (Appendix I).

Oligophagy coefficients. — Gramineae
1.000; Chloridoideae 0.874; Muhlenhergia
richardsonis 0.874. [Note: M. richardsonis
has been present in all communities where
decora occurs.]

Remarks and diagnosis. — Many individu-
als of decora have a rather distinctive habitus,
with very dark fore wing markings, especially
on the apical portions, and along the veins of
the forewing that contrast sharply with the
lighter pronotum and crown. Unfortunately,
paler specimens are not rare; the identity of
these specimens is often indicated by the ex-
tremely short crown length. Flexamia decora
and its close sister species, youngi, are appar-
ently restricted to Muhlenhergia richard-
sonis.

28. Flexamia youngi, n. sp.

Length of c? 3.7 (3.4-3.9) mm, 9 3.8
(3.7-4.2) mm; head width of 6 1.12 mm, 9
1.18 mm. Crown not produced; median
length of crown approximately 1.11 x head
width and 0.58 x interocular width (c? n = 16;
9 n = 20).

Color Stramineous. Crown with at most a
trace of basal or midlength markings. Prono-
tum with or without traces of wide, brown
stripes. Face with black interocular line con-
trasting with white lower face. Venter strami-
neous.

Male. — Pygofer (Fig. 38D) as in decora
but with posterior lobe less angulate, more

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Great Basin Naturalist Memoirs

No. 12

Fig. 38. Flexamia youngi, n. sp.: A, aedeagus and connective, lateral aspect; B, aedeagus and connective, dorsal
aspect; C, male plates and pygofers, ventral aspect; D, male pygofer, lateral aspect; E, right style, dorsal aspect; F,
female sternum VII; G, bases of first valvulae of female, dorsal aspect.

broadly rounded ventrally; plates (Fig. 38C)
extending approximately 2/3 length of pygo-
fer, fused basally for 2/3 length, connective in
lateral aspect (Fig. 38A) with dorsal keels
broad, approximately 1/3 height of dorsal
apodeme; apodemal processes sinuate in ven-
tral view, bell-shaped in caudal aspect; styles
as in other members of decora subgroup;
aedeagus (Fig. 38B) asymmetrical, shaft elon-
gate, apparently twisted 1/8 turn counter-
clockwise in caudal aspect, tapering gradually
in lateral aspect, slightly enlarged apically in
ventral aspect; three apical processes extend-
ing basad; unpaired process recurved, entire,
with median ventral groove almost 1/2 length
of shaft. Paired processes recurved, serrate on
dorsal edges, approximately 1/3 length of un-
paired process. Gonopore small, circular, and
apical.

Female. — Sternum VII (Fig. 38F) with
posterior margin produced medially, pro-

nounced median notch with infuscated spots
on either side. Ovipositor with base of first
valvula (Fig. 38G) recurved, appearing tear-
shaped and only slightly exceeding lateral
margins when viewed dorsally.

Types. — Holotype 6 : White Pine County,
Nevada, Baker, 10 August 1986, R. F. Whit-
comb (IPL 002590, Muhlenhergia richord-
sonis). Deposited USNM. Paratypes: 10 6,
119, same collection data; 1 9 , Mt. Wheeler,
White Pine County, Nevada, 10 August 1986,
8,500 ft, IPL 002569; 4 d , 4 9 , Spring Valley,
White Pine Countv, Nevada, 11 August 1986
(5,800 ft, R. F. Whitcomb, IPL 002596). De-
posited in BARC, CNC, KSU, and USNM.
Paratypes (KU) were also collected by R. H.
Beamer (1 9, Deeth, Elko Co., Nevada, 21
July 1947); and D. H. Lindsay (1 d, 3 9
Soldier Summit, Utah, 13 August 1936).

Remarks. — This species, apparently a
Great Basin endemic (Fig. 39), is closely

1988

Whitcomb, Hicks: Flexamia Revision

269

Fig. 39. Geographic distribution o( Flexamia decora (•), youngi (O), and modica (I
richardsonis (decora and youngi) and M. repens (modica).

and their hosts, Mtihlenbergia

related to decora but can be distinguished
from it by the shape of the aedeagus, which is
straight and twisted rather than curved and
symmetrical, and by the recurved, apical pro-
cesses. The host so far as known is (like de-
cora) Mtihlenbergia richardsonis. This spe-
cies is named in honor of D. A. Young, whose
classic monograph, with B. P. Beirne, estab-
lished the modern generic concept of Flex-
ainia.

29. Flexamia modica Beamer & Tuthill
Flexamia modica Beamer & Tuthill 1934; 3.

Important CHARACTERS. — Length of 6 3.4
(3.1-3.7) mm, of 9 3.8 (3.4-4.0) mm. Crown
moderately produced (1.26 x interocular
width, 0.62 X head width) (c? n = 20; 9 n =
20). Face ivory, with broad interocular band.
[6] Pygofer (Fig. 62JJ) as in decora. Plates
(Fig. 33F) short, about 2/3 length of pygofer,
broadly triangulate, widely divergent at
apices. Aedeagus (Fig. 31 M) asymmetrical,
gonopore triangular; unpaired ventral process
less than half length of shaft, base appearing
fiised to shaft in lateral aspect. [9] Sternum

VII with broad median projection; ovipositor
with base of each first valvula turned dorsad
through 90 degrees and bilobed, recurved
portion extending strongly laterad and ex-
ceeding lateral margin of valvula in dorsal as-
pect.

Geographic distribution. — High desert
plains of southern New Mexico and adjacent
counties of Trans-Pecos Texas and Arizona
(Fig. 39). It is abundant in the Sacramento
Mountains of southeastern New Mexico on its
host Muhlenbergia repens. It is much less
common in the Gila Mountains, where its host
is more patchy, and occurs rarely in the cen-
tral Guadalupe Mountains (Guadalupe Moun-
tains National Park, Culberson Co., Texas)
and northeastern Arizona (St. Johns, Apache
Co.).

Biology — Apparently a specialist oi Muh-
lenbergia repens. Our records show seven se-
ries from this host 16-18 June 1985 (R. F.
Whitcomb and K. A. Allred, approximately
440 S, 9, and immatures). M. repens is
closely related to M. richardsonis (Morden
1985, Morden and Hatch 1987). It is likely
that modica arose from the decora lineage via

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a host transfer from M. richardsonis.

Oligophagy coefficients. — Gramineae
1.000; Chloridoideae 1.000; Muhlenbergia
repens 1. 000 {n = 377).

30. Flexamia inflata (Osborn & Ball)

Deltocephalus inflatus Osborn & Ball 1897: 202.
Deltocephalus (Flexamia) inflatus, DeLong 1926: 3.5.
Flexamius inflatus, DeLong and Sleesman 1929: 84.

Important characters. — Length of 6 3.8
(3.1-4.4) mm, of 9 4.0 (3.6-4.6) mm. Crown
short (1.11 X interocular width; 0.58 x head
width) ((5 n = 26; 9 n ^ 26). Face strami-
neous, with or (more frequently) without a
basal, pale brown interocular band. [6]
Pygofer (Fig. 62GG) with upper portion of
posterior lobe strongly produced posteriorly.
Plates (Fig. 34A) short, about 1/2 length of
pygofer, apically divergent. Aedeagus (Fig.
31 E) asymmetrical; shaft elongate, slender,
gradually broadened apically; gonopore apical
on caudoventral surface; pair of short, re-
curved processes arising on each side of
gonopore, slender, unpaired ventral process
almost half length of shaft, slightly expanded
anteapically, acute at apex, which extends to
right of shaft, all processes with edges entire.
[9] Female sternum VII broadly, medially
produced, medially notched with traces of
teeth between notch and lateral margin;
ovipositor with base of each first valvula (Fig.
63T) curved dorsad, small projection on dorsal
edge extending dorsolaterad, entire curved
portion more heavily sclerotized and appear-
ing almost as separate sclerite in ventrolateral
aspect.

Geographic distribution. — This species
is distributed (Fig. 41) from Utah, Texas, and
New Mexico north to British Columbia and
Manitoba east to Pennsylvania.

Biology. — This species may be adapted to
northern mixed grasslands where pooid
grasses, Juncus spp. , and Muhlenbergia grow
in intermixed stands. In the Southwest, at
least, inflata is a specialist on Muhlenbergia
asperifolia ; occasional collections have also
been made on Distichlis spicata. We do not
have host records from the Northwest, but the
northern limit of the distribution of inflata
coincides roughly with that of Muhlenbergia
asperifolia. In Illinois it occurs on Juncus
tenuis. In Maryland it occurs on J uncus tenuis
and on the introduced grasses Cynodon dacty-

lon, Zoysiajaponica, and Eragrostis curvula.
This species has been reared on /. tenuis
(Whitcomb 1957). Populations occasionally
turn up for a time on Festuca or Poa but never
seem to persist. We have observed no mor-
phological indications of host races, although
it is possible that the Juncus populations may
be reproductively isolated from other popula-
tions. Rearing studies may be required to re-
solve the status of the biology oi inflata.

Oligophagy coefficients. — Prairie and
savanna: Gramineae 0.725, Juncaceae 0.275;
Chloridoideae 0.725, Muhlenbergia asperifo-
lia 0.562; Eragrostis curvida 0.066 (n = 379).
Other records: Manitoba (Woodside, 24 Au-
gust 1981, 16 d , 16 9 , Poa pratensis, K. G. A.
Hamilton, CNC).

Remarks and diagnosis. — Although defin-
itive identification of inflata requires exami-
nation of the genitalia, it is important to learn
to sort the majority of specimens of this often
abundant species on the basis of habitus
alone. The vast majority of specimens of in-
flata are very lightly pigmented; these can be
recognized by their light stramineous color in
combination with a black spot on the apical
forewing cell (Fig. 4A), by the lack of a spot in
the corium, and by their crown, which is one
of the shortest in the genus.

31. Flexamia beameri, n. sp.

Length of 6 3.2 (3.1-3.3) mm, 9 3.4
(3.3-3.4) mm; head width of 6 0.93 mm, 9
1.06 mm. Crown moderately produced; me-
dian length of crown 0.67 x head width and
1.39 X interocular width (d n = 6; 9 n = 2).

Color stramineous with dark markings on
corium and apical cell of forewings; face as in
inflata and texana; head with light markings,
pronotum with wide, faint stripes; venter and
legs with irregular, fuscous markings.

Male. — Pygofer (Fig. 40D) as in texana;
plates extending to approximately 2/3 length
of pygofer, fused basally for 3/4 length; con-
nective in lateral aspect (Fig. 40A) with dorsal
keels broad, approximately 2/3 height of dor-
sal apodeme; apodemal processes narrowly
campanulate in caudal aspect; style (Fig. 40E)
as in inflata and texana; aedeagus (Figs.
40A,B) slightly asymmetrical, shaft sigmoidal
in lateral view, ventral unpaired process not
quite in median plane of shaft and appearing
slightly longer than strongly recurved,
paired, lateral processes; all processes with
edges entire; gonopore circular and apical.

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Fig. 40. Flexamia heameri, n. sp.: A, aedeagus and connective, lateral aspect; B, aedeagus and connective, dorsal
aspect; C, male plates and pygofers, ventral aspect; D, male pygofer, lateral aspect; E, right style, dorsal aspect; F,
female sternum VII; G, bases of first valvulae of female, dorsal aspect.

Female. — Sternum VII (Fig. 40F) similar
or identical to texana, with median notched
projection adjoined on each side by serrulate,
infuscated projections; ovipositor with re-
curved portion of the bases of each first
valvula (Fig. 40G) narrowing to diverging
apices in dorsal aspect.

Types. — Holotype 6: Otter Lake, Oneida
County, New York, 25 June 1946, R. H.
Reamer. Deposited KU. Paratypes: 5 S and 2
9 , same locality, deposited KU and USNM.

Remarks. — This species, known from a sin-
gle collection at its type locality (Fig. 41), is a
sister to texana, a species also known from a
single collection made long ago. Flexamia
beameri can be readily separated from inflata
and texana by the shape and length of the
aedeagal processes. The aedeagal characters
oi beameri resemble those of texana, but the
processes of beam£ri are not twisted. Pre-
sumably, the ranges oibeameri and texana are

very different. In contrast to the hot climate of
south central Texas, the type locality of beam-
eri is located in one of the coolest regions of
New York (see Notes, Appendix III). Since we
have not yet encountered these species in the
field, their biology is a mystery. This species
has been named in honor of R. H. Reamer,
who described several interesting species of
the genus and who collected the holotype and
paratypes at a geographic site far from his
home base.

32. Flexamia texana Young & Reirne

Flexamia texana Young & Beirne 1958: 29.

Important characters. — Length of 6
3.1 (2.9-3.2) mm, of 9 3.2 (3.0-3.5)
mm. Crown not strongly produced
(1.30 X interocular width, 0.63 x head width)
(c? n = 8; 9 n = 2). [(?] Pygofer (Fig. 62HH)
with upper margin flat in lateral aspect, pos-
terior margin gradually rounded. Aedeagus

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Fig. 41. Geographic distribution oi Flexamia inflata (•), beameri (A), and tcxana (O).

(Fig. 31A) asymmetrical; unpaired ventral and
pair of lateral processes, all about equal in
length, arising near gonopore; ventral process
not in sagittal plane, lateral processes not in
symmetrical planes. [ 9 ] Sternum VII as for
group; ovipositor with base of each first
valvula (Fig. 63CC) curved dorsad through
about 90 degrees in lateroventral aspect, base
acute in lateral aspect, regularly convex in
dorsal aspect.

Geographic distribution. — Known only
from the type locality (Uvalde, Texas).

Biology. — The host oitexana is unknown.
Uvalde is situated just south of the southern
edge of the Edwards Plateau. We have sur-
veyed the main, dominant, warm-season
grasses of the Edwards Plateau searching for
this species. Although these searches turned
up a new species, collorum, they failed to
locate texana. The grasslands of the mesquite-
acacia savannah south of Uvalde have been
largely plowed for agricultural crops. Perhaps
E. D. Ball was afforded an opportunity to
collect in these grasslands before they were
sacrificed for agriculture. If this is the explana-
tion for the rarity of this species, we may
experience continuing difficulty in our search

for this elusive species. On the other hand,
discovery of the sister species heameri, whose
type locality suggests a marshy wetland, raises
the possibility that texana may turn up in
wetlands in south Texas, which were over-
looked in our searches.

XI. The imputans Group

The imputans group consists of a single spe-
cies, which is apparently a Muhlenbergia spe-
cialist in the northern prairie. This species
retains the plesiomorphic articulation be-
tween connective and aedeagus; it could oth-
erwise be regarded a close sister species to
areolata, with which it shares a distinctive
aedeagal morphology. These two species are
both greenish dorsally and have black faces.

33. Flcxamia imputans (Osborn & Ball)

Deltoccphditis i)nput(in.s O.slwrn 6c Ball I89S: 75.
Deltoceplialti.s (Ficxinnia) iiti})utiius, DeLong 1926: 29.
Flexainius [.sic] imputans, DeLong and Caldwell 1937; 27.

Important CHARACTERS. — Length of 6 3.4
(3.1-3.7) mm, of 9 3.7 (3.3-3.9) mm. Crown
moderatelv produced (1.33 x interocular
width; 0.62 X head width) (d n = 10; 9 n =

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273

Fig. 42. Geographic distribution oi Flexamia imputans and a probable host, Muhlenbergia cuspidata.

14). Face usually entirely black but occasion-
ally with oblique, pale streak on each gena.
[6] Genitalia (Fig. 31B) similar to those of
areolata but with hind margin of pygofer (Fig.
62V) convex and with distinct joint between
connective and aedeagus. [9] Sternum VII
(Fig. lOP) with hind margin broadly, slightly
convexly produced with slight median notch;
ovipositor with bases of first valvulae (Fig.
63II) not recurved.

Geographic distribution. — Northern
Great Plains from the Dakotas south to Kan-
sas, east to Iowa and Wisconsin (Fig. 42).

Biology. — This species is apparently a
Muhlenbergia specialist and may occur
largely on Muhlenbergia cuspidata. It is one
of the rarest of the Flexamia species and,
given the extensive turnover of its original
upland prairie habitat to croplands, may be-
come increasingly rare.

Remarks and diagnosis. — This species is
unique in its lack of markings on head and
pronotum, in combination with its black face.

XII. The areolata Group
This group consists of a single species.
34. Flexarnia areolata (Ball)

Deltocephalus areolatus Ball 1899: 188.
Deltocephahis (Flexamia) areolatus, DeLong 1926: 25.
Flexamia areolatus, DeLong and Sleesman 1929: 84.

Important gharacters. — Length of (5 2.8
(2.6-3.2) mm, of 9 3.6 (3.2-4.0) mm. Crown
strongly produced (1.77 x interocular width;
0.75 X head width) (c? n - 20; 9 n = 22). Face
entirely black. [6] Pygofer (Fig. 62Z) with
dorsal portion of posterior lobe produced cau-
dodorsad, posterior margin oblique and
slightly concave. Plates (Fig. 34G) elongate,
gradually narrowed from base to apex, fused
basally for about 1/3 length. Aedeagus and
connective fused. Aedeagus (Fig. 31C) asym-
metrical; three tapering apical processes, two
short, with serrate edges, extending basad
along right side of shaft; third process extend-
ing basad for about half the length of right side
of shaft, minute projections in middle,
broader than other two processes, bearing
gonopore as elongate slit through almost en-
tire length. [6] Sternum VII (Fig. lOQ) with
posterior margin concave on each side of api-
cally notched, median, convex projection;
ovipositor with first valvulae not curved dor-
sad at bases.

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Fig. 43. Geographic distribution ofFlexamia areolata and its host, Eragrostis spectabilis.

Geographic distribution — This species
occurs from Connecticut to Wisconsin, south
to Florida and Texas, covering much of the
geographic range of its host (Fig. 43). One of
the three specimens from the original cotype
series is labeled "Phoenix, Arizona, May,
1897." One specimen in the CNC is labeled
"San Diego, Calif., 19-6-71, GL1251, H. H.
Ross." Nevertheless, occurrence of areolata
in the Southwest requires confirmation.

Biology. — We consider areolata to be a
specialist of Eragostis spectabilis. DeLong
(1949) believed that the host was "Panicum or
a closely related genus." Panicum capillare,
an annual that commonly occurs with E.
spectabilis in sandy habitats, has a superfi-
cially similar inflorescence characterized by
wide panicles that redden when mature. As if
this were not confusing enough, Leptoloma
cognatum, a representative of a genus that is
indeed closely related to Panicum, not only
occurs with the above two plant species in the
same sandy habitats, but also has low, wide
panicles that redden when mature. Unsure
whether we may have overlooked compo-
nents of this confusing plant association, we
have returned to several communities where

areolata occurs and have found E. spectabilis
growing in the absence of the other two
grasses. Furthermore, we have collected this
species in the Nebraska Sand Hills on E.
spectabilis, northwest of the range of Lep-
toloma cognatum. Osborn (1928) recorded
areolata from Eragrostis pectinacea in North
Carolina. Because E. pectinacea is an annual,
this report requires confirmation.

Remarks and diagnosis. — The habitus of
areolata is unique.

XIV. The prairiana Group

The prairiana group is named for its
most plesiomorphic member, F. prairiana,
a widespread resident of tall-grass prairie
and southwestern semiarid grasslands. The
group consists of 10 species that share a simi-
lar general facies. All members of the group
are medium-sized and brown but lack distinc-
tive markings. Definitive identification re-
quires examination of genitalia; the aedeagus
and connective are fused, a condition we de-
fine as synapomorphic. In prairiana a trace of
the plesiomorphic articulation remains and
constitutes an autapomorphy defining that
species. Species of this complex are largely

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275

inhabitants of prairie, savanna, or forest
glades. Members of the group speciahze
largely on andropogonoid grasses {Andro-
pogon, Schizachyriiwi, and Botliriochloa
spp.). Exceptions are (1) atlantica, which is
often associated with switchgrass {Panicwn
virgatU7n) or, in the Southwest, chloridoid
grasses such as Sporoboliis spp.; and (2)
producta, a southern coastal species that ap-
pears to be associated with chloridoid grasses.
Both atlantica and producta regularly colo-
nize an introduced chloridoid grass, Cynodon
dactylon.

There are two sets of closely related sister
species. One is sandersi-delongi. Ross and
Cooley (1969) separated delongi from sandersi
on the basis of the greater divergence of its
apical, paired processes and lesser appression
of the unpaired process to the aedeagal shaft
(Figs. 44B,C). Hamilton and Ross separated
satilla from clayi on the basis of a narrower
aedeagal shaft (Figs. 44H,J). The distinctions
between the species of these two sets are fine;
some workers would consider the differences
to represent infraspecific geographic varia-
tion. However, we have retained satilla and
delongi because, in each case, they can be
unambiguously identified, and a geographic
and historical basis can be presented to ex-
plain both their divergence and maintenance
of reproductive isolation. In the case of de-
longi, which probably specializes on Schiz-
achyriiim scoparium, the geographic range
(Fig. 50) was almost entirely glaciated during
the most recent glacial maximum. In contrast,
sandersi occupies a region that was never
glaciated, where it apparently specializes
largely on Andropogon virginicus. These cir-
cumstances provide a hypothetical scenario
for speciation and an explanation of ecological
factors that may now enforce isolation be-
tween the two species. Similarly, satilla, de-
scribed by Hamilton and Ross (1975) from
south Georgia, is probably a Gulf Coast spe-
cies (Fig. 51) that is at least partially sympatric
with producta. Discovery of its sister species,
clayi, as far north as Maine suggests that dif-
ferentiation of these species, hke that of
sandersi and delongi, may be a postglaciation
event that is phenologically enforced by con-
temporary climate. As closely related as clayi
and satilla are, the sister set that they com-
prise is also very closely related to graminea.
A more distant pair of sister species is that of

prairiana and reflexa. The most significant
apomorphy shared by these sister species is
the dorsal position of the gonopore (Figs.
44D,G). These species also share a common
general facies and cannot be distinguished on
the basis of external characters. Each of these
sister species has a variable aedeagal morphol-
ogy, and we found that careful study was re-
(juired to separate them. Both species are
residents of tall-grass prairie, where they oc-
cur on Andropogon and Schizachyrium spe-
cies. Other species of the prairiana group are
more distantly related.

Description of prairiana Group

Medium-sized. Length of 6 3.0-4.2 mm,
of 9 3.0-4.6 mm. All species with similar
general facies. Brown, or in darkest speci-
mens, black on face or venter. Vertex moder-
ately produced. Crown often stramineous
with conspicuous transverse line at mid-
length. Male plates elongate, in some species
notched apically. Aedeagus and connective
fused, or with at most (in prairiana) a faint
trace of the plesiomorphic articulation. Ae-
deagus with paired, apical, serrate processes
and an unpaired, lateral or dorsal process par-
tially fused to shaft in some species, or in
others entirely fused or absent. Aedeagus
asymmetrical in all but one species (dakota).
Gonopore in primitive species oval and an-
teapical, in more advanced species located on
the unpaired process or, in the most highly
specialized species, replacing the unpaired
process as a spiral groove originating dorsally
on the shaft. Female sternum VII produced
posteriorly at middle in a convex, apically
notched protuberance.

Key to Males of the prairiana Group
1. Aedeagus with 3 processes on shaft 2

— Aedeagus with 2 processes on shaft 6

2(1). Pygofer with posterior lobe strongly expanded

on ventral margin (Fig. 5D) . . . producta (Walker)

— Pygofer with ventral margin of posterior lobe
not strongly expanded 3

3(2). Gonopore dorsal, anteapical, located on a pro-
cess or at the base of one (Figs. 44D,G) 4

— Gonopore on dorsolateral surface of shaft at base

of retrorse process 5

4(3). Gonopore near apex of aedeagal shaft, at base of

a process (Fig. 44G) prairiana DeLong

— Gonopore located on a process (Fig. 44D) ....
reflexa (Osborn & Ball)

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Fig. 44. Aedeagal apices of the Flexamia prairiana group: A, atlantica- B, sandersi- C, delongi- D, reflexa; E,
producta; F, graminea; G, prairiana; H, c/flyi, I, dakota; J, satilla. Aspects: AD, anterodorsal; CD, caudodorsal; CE,
cephalic; CV, caudoventral; DS, dorsal; LT, lateral; VN, ventral. Redrawn from Young and Beirne (1958) and Hamilton
and Ross (1975).

5(3). Apical processes of aedeagus more divergent,
each with 6-9 teeth; when viewed in a position
with apical processes horizontal, a space is visi-
ble between the base of the gonopore extension
and the shaft (Fig. 44C) . . . delongi Ross & Cooley

Apical process of aedeagus less divergent, each
with 3-5 teeth; when viewed as above, no space
is visible between the base of the gonopore ex-
tension and the shaft (Fig. 44B)

saiidersi (Osborn)

6(1). Gonopore spiral 7

— Gonopore not spiral 8

7(6). Aedeagal appendages of aedeagus without
prominent, anteapical processes; aedeagal shaft
usually not strongly expanded at base of
gonopore (Fig. 44F) graminea (DeLong)

— Apodemal appendages of aedeagus each with
prominent, anteapical protuberance directed
mesad; aedeagal shiift strongly expanded at base

of gonopore (Fig. 44H) 9

8(6). Aedeagal shaft with apical process on each side;
gonopore anteapical on anterodorsal surface
(Fig. 441) dakota Young & Beirne

— Aedeagal shaft with apical processes arising on

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Whitcomb, Hicks: Flexamia Revision

277

one side; gonopore anteapical on lateral surface
of'shaft (Fig. 44A) atlantica DeLong

9(7). Shaft of aedeagus comparatively robust (Fig.

44H); north Georgia to Maine

claiji Young & Beirne

— Shaft of aedeagus slender (Fig. 44J); Gulf Coast

(Florida, south Georgia, Mississippi)

satilla Hamilton & Ross

Key to Females of the prairiana Group

1. Ovipositor with base of each first valviila re-
curved dorsally in dorsal aspect 2

— Ovipositor with base of each first valvula not
recurved dorsally 5

2(1). Recurved portion of each first valvula parallel to
anteriormost border of each valvula in dorsal
aspect, appearing as rim above the valvula (Fig.
63LL) 3

— Recurved portion appearing wider in dorsal as-
pect prairiana DeLong

3(2). Anteriormost border of each first valvula ap-
pearing somewhat angular in dorsal aspect, re-
curved portion very narrow and not produced
dorsad (Fig. 63LL) graminea DeLong

— Anteriormost border of each first valvula
rounded, recurved portion much broader and
produced dorsad (Fig. 63GG) 4

4(3). Range north Georgia to Maine

clayi Young & Beirne

— Range Gulf Coast satilla Hamilton & Ross

5(1). Bases of first valvulae obliquely truncate, valvu-
lae apparently membranous basally (Fig. 63HH)
reflexa (Osborn & Ball)

— Bases of first valvulae convex basally 6

6(5). First valvulae broadest near bases in dorsal as-
pect (Fig. 63EE,KK) 7

— First valvulae broadest more posteriorly in dor-
sal aspect 8

7(6). Face often with brown interocular line contrast-
ing with pale color of remainder; if without line,
then irregularly darkened; Southeast (Fig. 49)
producta (Walker)

— Face tan to dark brown with alternating sinuate,

dark and light lines; dry prairie (Fig. 49)

dakota Young & Beirne

8(6). Face with black interocular line, or at least with
this area black in addition to other black marks
on face atlantica DeLong

— Face brown or tan; interocular area never black 9

9(8). Range southern Illinois southeastward

sandersi (Osborn)

— Range northern Illinois northward to Ontario,

northwest to North Dakota

delongi Ross & Cooley

35. Flexamia prairiana DeLong
Flexamia prairiana DeLong 1937: 32.

Important CHARACTERS. — Length of 6 3.6

(3.2-3.9) mm, of 9 3.9 (3.4-4.6). Crown
moderately produced (0.60 x head width, L40
X interocular width) (d n 73; 9 n 32).
Face (Fig. 3D) usually pale in lower half, with
broad, black interocular band; in darkly
marked specimens almost completely black
(Fig. 3C). Habitus characteristic of group.
Male plates (Fig. 45A) elongate, unnotched.
[6] Pygofer (Fig. 62W) with upper part of
hind margin produced posteriorly. Aedeagus
(Figs. 6F, 44G) asymmetrical, shaft slender,
elongate, gradually broadened apically; gono-
pore oval, anteapical on anterodorsal surface.
Three retrorse apical processes diverging
slightly from shaft, about one-third length of
shaft. Unpaired process arising basad of
gonopore, margins entire with dorsal, longi-
tudinal groove on apical half; pair of distad,
ventral, lateral processes, serrate on ventral
margin. [ 9 ] Sternum VII as for group; ovipos-
itor with each first valvula (Fig. 63 JJ) recurved
slightly dorsad at base.

Geographic distribution. — Widely dis-
tributed (Fig. 46) from Manitoba to Colorado,
Arizona, and Mexico, east to Missouri and
Illinois. Two females from Michoacan, Mex-
ico (21 October 1981, M. W. Nielson), have
been tentatively referred to prairiana. A sin-
gle male (Baker 2361, USNM) is recorded
from Auburn, Alabama, 10 September 1897.

Biology. — Southwestern populations usu-
ally occur on Bothriochloa species. In the
tall-grass prairie the usual hosts are Schiz-
achyrium scoparium and Andropogon ger-
ardii. Most reported hosts are andropogonoid
grasses, but in tall-grass prairie prairiana may
move short distances to more suitable hosts in
drought periods or in late summer. Thus,
oligophagy coefficients calculated for prairi-
ana tend to be among the lowest for all Flex-
amia species (Whitcomb et al. 1988). The
green coloration of live specimens of prairi-
ana fades to orange upon drying. In midsum-
mer the intensity of coloration varies greatly.
In the fall, coloration may be very dark; un-
cleared, late-season specimens have often
been misidentified.

Oligophagy coefficients. — Tall-grass and
mixed-grass prairies north of Texas: Grami-
neae 0.922; Panicoideae 0.730; Schiza-
chyrium scoparium 0.438; Andropogon ger-
ardii0.1A\{n -527).

Remarks and Diagnosis. — Because this
species is one of the most abundant of Flex-
amia species, it is important to recognize it

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Fig. 45. Male plates and pygofer.s of the Flexamia imiiriana group, ventral aspect: A, prairiana; B, reflexa; C,
atlantica; D, producta; E, dakota- F, sandersi; G, delun^i; H, graminea; I, clayi.

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Whitcomb, Hicks: Flexamia Revision

279

Fig. 46. Geographic distribution of Flexamia prairiana.

(tentatively) without dissection. The prairi-
ana face type is almost always distinguishable
from the faces ofgraminea, sandersi, delongi,
or dakota, which may occur with it in Andro-
pogon prairies. Dissected male specimens re-
veal an aedeagus that might be confused only
with that of reflexa. Female specimens may
prove difficult to identify because the bases of
the first valvulae vary geographically. In
southwestern populations this structure is
rarely sclerotized, as illustrated by Young and
Beirne (1958).

36. Flexamia reflexa (Osborn & Ball)

Deltocephalus reflexus Osborn & Ball 1897: 203.
Deltocephalus (Flexamia) reflexus, DeLong 1926: 28.
Flexamius reflexus, DeLong and Caldwell 1937: 27.

Important CHARACTERS. — Length of S 3.5

(3.1-4.0) mm, of 9 3.7 (3.2-4.2) mm. Crown
moderately produced (1.49 x interocular
width; 0.68 X head width) (c? n = 20; 9 n =
15). Face variable, often pale on lower half
with broad interocular band; band may be
obscured in dark specimens. [6] Pygofer
(Fig. 62X) with posterior lobe strongly pro-
duced, truncate apically. Plates (Fig. 45B)
elongate, without notches or lateral projec-
tions. Connective in lateral aspect with dorsal
keels extending almost as far dorsad as aedea-
gal apodeme. Aedeagus (Fig. 44D) asymmet-
rical, shaft elongate, gradually tapered; gono-
pore anteapical on anterodorsal surface, asso-
ciated with short, apically grooved, unpaired
process. Paired apical processes serrate along
ventral edge, extending basad less than half
length of shaft in lateral aspect. [ 9 ] Ovipositor

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Fig. 47. Geographic distribution of Flexamia reflexa and a host, Andropogon gerardii.

with first valvulae membranous basally, not
curved dorsad; sclerotized portion obliquely
truncate in dorsal aspect.

Geographic distribution. — Tall-grass
prairie (Cwikla and Blocker 1981), savanna,
and glades and prairie mosaic of deciduous
forest, Kansas and Iowa to Texas, Ontario,
Maryland, and Florida (Fig. 47).

Biology. — This species is associated with
Andropogon gerardii in the East (Maryland
and West Virginia) and may be associated with
that host in the tall-grass prairie. However, in
Texas it occurs in mixed-prairie associations in
which big bluestem is a minor component.

Diagnosis and remarks. — Given the pau-
city of specimens of reflexa in collections, it
is remarkable that it was the second member
of the prairiana group to be described and
that it is the type species of the genus. Almost
all uncleared specimens "identified" as this
species in museum drawers turn out to be
other prairiana group species. The abundant
prairiana, which occurs through much of its
range on httle bluestem {Schizachyrium sco-
parium), is much more apt to be encountered
in the prairie. Since we have found no reliable
external characters for sorting reflexa from

prairiana, a search for this species involves
sifting through numerous prairiana speci-
mens. The position of the gonopore (some
distance from the aedeagal apex, situated
basally on a short, appressed, unpaired pro-
cess) contrasts sharply with the simpler, oval,
anteapical gonopore oi prairiana . Some spec-
imens of atlantica and producta may also re-
semble reflexa in general facies, but their gen-
italia bear little resemblance to those of
reflexa.

37. Flexamia atlantica (DeLong)

Deltocephahis (Flexamia) atlanticus DeLong 1926: 29.
Deltocephalus atlanticus, DeLong and Caldwell 1937; 27.

Important characters. — Length of (5 3.7
(3.2-4.2) mm, of 9 4.1 (3.6-4.5) mm. Crown
moderately produced (1.31 x interocular
width; 0.59 X head width) (d n 20; 9 n =
20). Face black in upper half, often forming
interocular band contrasting with pale lower
half. [6] Pygofer (Fig. 62Y) with posterior
lobe narrow, without ventral lobe. Plates
(Fig. 45C) elongate, notched apically. Aedea-
gus (Fig. 44A) asymmetrical, shaft nearly uni-
form in width, gonopore simple, anteapical on

1988

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281

Fig. 48. Geographic distribution of Flexamia atlantica and a host, Panicum virgatum.

the right side. Pair of elongate, retrorse, ser-
rate, apical processes extending basad approx-
imately half length of shaft. [9] Sternum VII
as for group; ovipositor with base of each first
valvula (Fig. 63DD) not curved dorsad.

Geographic distribution. — Prairie from
Manitoba, North Dakota, Wyoming, and New
Mexico east to Illinois (Fig. 48). Also along the
Atlantic coastal prairie of the Northeast, and
very patchily in the Southeast.

Biology. — In tall-grass prairie atlantica is
usually associated with switchgrass {Panicum
virgatum). This grass, a dominant in the
northeastern coastal prairie, probably ac-
counts for the occurrence of atlantica there.
In mixed-prairie and southwestern grass-
lands, atlantica occurs on native Sporobolus
spp. and is the major Flexamia colonist in
the extensive stands of two exotic chloridoid
grasses (weeping love grass [Eragrostis cur-
vula ] and bermudagrass [Cynodon dactijlon ])
in Texas and Oklahoma.

Oligophagy coefficients. — Texas and
Oklahoma: Gramineae 0.994; Chloridoideae
1.000; Eragrostis curvula 0.243; Cynodon
dactylon 0.633 (n = 161). Kansas, Missouri,
and Nebraska: Gramineae 1.000; Panicoideae

0.876; Panicum virgatum 0.783; Sporobolus
spp. 0.124 (n = 177).

Diagnosis and remarks. — Flexamia at-
lantica is the only pale-faced Flexamia prairie
species with apically notched plates. Also,
individuals of atlantica tend to be larger than
those of reflexa or prairiana. In the South-
east, atlantica may occur with producta,
which tends to be smaller, and which usually
has a much more produced crown. Although
these characters permit tentative recognition,
definitive recognition is readily accomplished
by examining the aedeagal morphology,
which is unique.

38. Flexamia producta (Walker)

lassus productus Walker 1851: 891.
Deltocephalus visendus Crumb 1915: 189.
Deltocephahis (Flexamia) productus, DeLong 1926: 43.
Flexamia producta, DeLong and Knull 1945: 35.

Important CHARACTERS. — Length of c? 3.2
(3.0-3.5) mm, of 9 3.5 (3.2-3.9) mm. Crown
usually conspicuously produced (1.62 x inter-
ocular width; 0.75 x head width) (cJ n = 20; 9
n = 20). Face variable but usually with clearly
delimited interocular band; band occasionally
obscured in darker specimens. [6] Plates

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Fig. 49. Geographic distribution of Flexamia dakota {•) and producta (■) and the host of dakota, Scizachyrium
scoparium, which does not occur on the Atlantic coastal plain.

(Fig. 45D) elongate, notched at apex. Pygofer
(Fig. 62AA) with posterior lobe produced and
rounded apically, ventral portion extending
mesad as pronounced lobe, serrate along mar-
gin, the two lobes often overlapping in ventral
aspect (but not necessarily in prepared speci-
mens). Aedeagus (Fig. 44E) asymmetrical api-
cally; shaft elongate, slender, slightly swollen
anteapically; gonopore irregularly oval, on left
side of dorsal surface of shaft; pair of processes
arising asymmetrically at apex of shaft, ex-
tending basad almost half of shaft length, each
process laterally compressed and conspicu-
ously toothed along ventral edge; with short,
small, curved process arising near proximal
portion of gonopore, extending sinistrad. [ 9 ]
Female sternum VII as for group, ovipositor
with bases of first valvulae (Fig. 63EE) not
curved dorsad, broad basally in dorsal aspect.

Geographic distribution.— Gulf Coast
and Atlantic seaboard from Mississippi to
Maryland (Fig. 49).

Biology. — This species has been collected
commonly in coastal prairies, but there are
few host records. We have recorded it from
several exotic chloridoid grasses (e.g., Zoysia

japonica and Cynodon dactylon) in Maryland
but have little field experience in the coastal
prairies where it is most abundant. We note
that Sporobolus virginicus is a common domi-
nant in southeastern coastal prairies; this chlo-
ridoid grass should be examined as a possible
natural host. On the other hand, there are
indications that cicadellid host specialization
decreases in mesic and wet prairies (Whit-
comb et al. 1987, 1988). Given its occurrence
in wet southeastern grasslands, producta may
turn out to be a habitat specialist.

Diagnosis and remarks,— The produced
crown of this species affords a tentative indica-
tion of its identity. Unfortunately, sotilla and
sandersi, which occur with producta, also
have produced crowns that approach the
length/width ratios oi producta. Although the
latter species are similar to producta in most
aspects, their faces are without conspicuous
interocular bands. It is therefore possible to
sort, from an assemblage of southeastern Flex-
amia, a large subset of producta specimens
that have either an extremely produced crown
or a distinct interocular band (or both). Unfor-
tunately, this will leave a mixture of atypical

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Whitcomb, Hicks: Flexamia Revision

283

producta specimens and specimens of
sandersi and satilla, or perhaps even clmji.
The aedeagal characters of producta readily
diagnose the species.

39. Flexamia dakota Young & Beirne
Flexamia dakota Young & Beirne 19.58: 44.

Important CHARACTERS. — Length of 6 3.0
(2.8-3.6) mm, of 9 3.3 (3.0-3.6) mm. Crown
moderately produced (1.40 x interocular
width; 0.64 X head width) {6 n = 21; 9 n -
18). Face brown to tan, with parallel lines. [ 6 ]
Pygofer (Fig. 62BB) with upper portion of
posterior lobe produced posteriorly. Plates
(Fig. 45E) with apical third narrowed. Aedea-
gus (Fig. 441) symmetrical; shaft elongate,
slender, narrower near midlength than at
base or apex. Gonopore anteapical on
anterodorsal surface; pair of recurved pro-
cesses extending basad amost half length of
shaft; each process serrate on ventral margin.
[ 9 ] Sternum VII as for group. Ovipositor with
bases of first valvulae broadest near base (Fig.
63KK).

Geographic distribution. — Semiarid
Great Plains of western North and South Da-
kota, eastern Wyoming, Nebraska Sand Hills
to Oklahoma, and the Edwards Plateau of
Texas (Fig. 49). A presumably disjunct popu-
lation occurs in the Loess Hills of western
Iowa. This species has recently been recorded
from Mexico (2 6 , 42 km S Piedras Negras,
Coahuila, 13 October 1987, A. L. Hicks and
J. E. Lowry).

Biology. — This species is apparently a spe-
cialist on little bluestem (Schizachyrium sco-
parium) in dry prairie.

Oligophagy coefficient, — Gramineae
1.000; Panicoideae 0.990; Schizachyrium sco -
pariwm 0.990 (n - 209).

Diagnosis and remarks. — This species of-
ten occurs in the absence of other specialists of
andropogonoid grasses. However, in the Ed-
wards Plateau of Texas, dakota occurs with
graminea (which it closely resembles in gen-
eral facies), prairiana, and, rarely, reflexa.
Although it can be tentatively sorted from
the latter two species on the basis of face
pattern, we have found no reliable external
character to sort it from graminea. The sym-
metrical aedeagus of dakota, which has only
paired apical processes, is unique. Young and
Beirne (1958) described only males; Lowry
and Blocker (1987) described the female.

40. Flexamia sandersi (Osborn)

Dritoci'pftalus sandersi Oshorn 1907: 164.
Deltocephaliis (Flexamia) sandersi, DeLong 1926: 27.
Flexamia sandersi. DeLong and Sleesman 1929: 83.
Flexamius bidentata DeLong 1935: 155.

Important characters. — Length of d 3.2
(2.9-3.5) mm, of 9 3.4 (3.2-3.9) mm. Crown
produced (1.48 x interocular width; 0.69 x
head width) (d n - 11; 9 n - 10). Face (Fig.
3H) varying from dark above and shading to
paler apically to brown or black throughout, in
former case with dark area shading gradually
along lower margin; no interocular band, [c?]
Pygofer (Fig. 62CC) truncate on dorsal por-
tion of posterior margin. Plates (Fig. 45F)
elongate, apically notched, in some speci-
mens bidentate. Aedeagus (Fig. 44B) asym-
metrical; shaft elongate, gradually tapered;
gonopore on dorsolateral surface of shaft at
base of retrorse, unpaired process, which is
closely appressed to the shaft, has one
minutely serrate margin, and extends basad to
midlength of shaft; pair of apical, retrorse pro-
cesses, dorsal slightly longer than ventral,
each acute apically and with distinctly serrate
margin. [ 9 ] Sternum VII as for group; ovipos-
itor with base of each first valvula (Fig. 63FF)
not curved dorsad.

Geographic distribution. — Eastern Kan-
sas to Louisiana, east to southern Illinois,
Massachusetts, and South Carolina (Fig. 50).

Biology. — The principal host appears to be
broomsedge (Andropogon virginicus), but
this conclusion requires further study.

Diagnosis and remarks. — Throughout
much of its range, sandersi occurs with clayi.
These species usually have been collected
in the summer before their host grasses
have flowered. Many host records are listed
simply as "Andropogon sp. ' In the Maryland
Piedmont sandersi occurs abundantly on
broomsedge in the absence of clayi; in Mary-
land we have found the latter species only in
the Appalachians. We have found no reliable
external character that separates sandersi
from clayi, satilla, graminea, or delongi.
However, the characters of the aedeagus de-
scribed above and in the key readily separate
sandersi from all other prairiana group spe-
cies, with the exception of delongi. We have
discussed the relationship of these very
closely related sister species in the corre-
sponding section under delongi.

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Great Basin Naturalist Memoirs

No. 12

Fig. 50. Geographic distribution of Flexamia sandersi (I
Andropogon virginicus.

and delongi (•) and the presumed major host oi sandersi.

41. Flexamia delongi Ross & Cooley

Flexamia delongi Ross & Cooley 1969; 246.

Flexamia sandersi. Young and Beirne 1958: 164. (In part.)

Important CHARACTERS. — Length of d 3.7
(3.4-4.1) mm, of 9 3.8 (3.4-4.4) mm. Crown
variably produced (1.34 x interocular width;
0.65 X head width) (c? n = 17; 9 n = 21). Face
varying from dark above, shading to paler api-
cally to black or brown throughout, dark area
often fading gradually along lower margin,
interocular band absent. [6] Pygofer and
plates as in sandersi. Aedeagus (Fig. 44C)
asymmetrical; shaft elongate, not cylindrical,
gradually tapered; gonopore on dorsolateral
surface at base of retrorse process, which ex-
tends basad to midlength of shaft; pair of lat-
eral, apical, retrorse processes extending
more than one-third but less than one-half
length of shaft, each with a distinct, coarsely
serrate margin and with very acute apex. [ 9 ]
Sternum VII as for group; ovipositor with base
of each first valvula as for sandersi.

Geographic distribution. — Northern In-
diana and Illinois, Wisconsin, and Ontario,
west to South Dakota, Saskatchewan, and
Manitoba (Fig. 50).

Biology. — This species is presumably a
specialist of little bluestem, Schizachyrium
scoparium.

Diagnosis and remarks. — The pair of sis-
ter species consisting of delongi and sandersi
is (with the exception of satilla and clayi) the
most closely related of all Flexamia sister
pairs. Ross and Cooley (1969) acknowledged
this closeness, pointing out that delongi pre-
viously had been considered a variant of
sandersi. Ross and Cooley examined material
from the East but did not examine specimens
from Canada or from the Dakotas. Thus, their
interpretation of delongi as a geographic re-
placement for sandersi made the identity of
the northern prairie material a crucial test.
We examined specimens from these regions
from the USNM and CNC and found them to
be referable to delongi. We retain herein the
concept of delongi, because it can be de-
fended on morphological, biological, and his-
torical grounds. The species occurs in a region
that was largely or completely glaciated dur-
ing the Wisconsinan. The principal (or sole)
host of sandersi {Andropogon virginicus)
does not occur in this region. Thus, san-
dersi and delongi appear to utilize different

1988

Whitcomb, Hicks: Flexamia Revision

285

hosts, whose phenological asynchrony may
reinforce reproductive isolation. F'inally, al-
though genitalic differences between delongi
and sandersi are slight, they are consistent.
The apical, aedeagal processes of delongi are
more divergent than those of sandersi; each
process has 6-9 teeth. Further, the gonopore
extension is more divergent from the shaft
than in sandersi.

In the northern prairies delongi may occur
on little bluestem with graminea, prairiana,
or dakota, but its aedeagal structure readily
separates it from these species.

42. Flexamia graminea (DeLong)

Deltocephalus (Flexamia) graniineus DeLong 1926; 30.
Flexamius graminetts, DeLong and Caldwell 1937: 27.

Important CHARACTERS. — Length of 6 3.6
(3.3-4.0)mm.,of 9 3.7(3.4-4.1) mm. Crown
moderately produced (1.40 x interocular
width; 0.68 X head width) (c? n - 25; 9 n -
20). Face variable, from entirely stramineous
to stramineous with clypeus darkened, from
tan to almost black with pale arcs; interocular
band absent. [S] Pygofer (Fig. 62EE) with
posterior lobe not well differentiated ven-
trally, posterodorsal portion produced cau-
dodorsad and truncate apically. Plates (Fig.
45II) divergent at apex, often appearing to be
notched. Aedeagus (Fig. 44F) asymmetrical;
shaft elongate, slender, twisted in apical half;
gonopore consisting of a spiral groove origi-
nating at shaft midlength; pair of asymmetri-
cal, retrorse, apical processes, with one
minutely serrate edge. [ 9 ] Sternum VII as for
group. Ovipositor with base of each first
valvula (Fig. 63LL) angular in dorsal aspect,
narrowly curved dorsad anterolaterally, re-
curved portion extending only slightly dor-
sad.

Geographical distribution. — Tall-grass
prairie of North Dakota, Kansas to Blackland
Prairie, and south Texas east to Missouri and
Illinois (Fig. 51). A specimen from the Ball
collection is labeled "DC." If this is an abbre-
viation for "District of Columbia," and if the
hastily scrawled, penciled label is correct,
then it would be necessary to explain a widely
disjunct distribution for graminea. Recently,
little bluestem and several unusual prairie
forb species have been found in a band of
savannalike habitat along the north bank of
the Potomac River between Great Falls Park
(Maryland) and the District of Columbia

(D. Boone, personal communication). The
possibility that graminea (and perhaps other
prairie insects) may have colonized such a
habitat cannot be completely discounted.

Biology. — This species appears to be a spe-
cialist of little bluestem (Schizachyrium sco-
pariiwi), but in late summer it may take
refuge on other hosts.

Oligophagy coefficient. — Schizachyri-
um scoparium 0.91.

Remarks and diagnosis. — Throughout
most of the tall-grass prairie, graminea occurs
regularly with prairiana on little bluestem. It
can be distinguished from prairiana by its
consistent lack of an interocular band and by
its notched plates. However, through por-
tions of its range it may also occur with
sandersi, dakota, or delongi, or in mixed-
prairie habitats with atlantica or reflexa. The
structure of its aedeagus separates it readily
from these species, but it is most closely re-
lated to clayi and satilla. For differential diag-
nosis of these three species, see the corre-
sponding section under clayi.

43. Flexamia clayi Young & Beirne

Flexamia clatji. Young and Beirne 1958: 64.

Important characters. — Length of 6 3.4
(3.3-4.0) mm, of 9 3.7 (3.1-3.8) mm. Crown
variably produced (1.44 x interocular width;
0.71 X head width) (c? n = 20; 9 n = 20). Face
brown to black above, usually shading gradu-
ally to paler below. [6] Pygofer (Fig. 62DD)
with posterior lobe more distinct than in
graminea. Plates (Fig. 451) divergent apically,
outer margins widened at midlength. Aedea-
gus (Fig. 44H) much as in graminea but with
shaft usually much more expanded at base of
gonopore. [9] Sternum VII as for group;
ovipositor with base of each first valvula
(Fig. 63GG) rounded, with recurved portion
broader than in graminea.

Geographic distribution. — Appalachian
region from Maine to north Georgia, west to
Illinois (Fig. 51).

Biology. — Some records for clayi are for
broomsedge {Andropogon virginicus); others
are for little bluestem {Schizachyrium scopar-
ium). Many are for "Andropogon." Since
broomsedge does not occur in Maine, at least
some of the records for little bluestem must be
correct. Many records, including our own, are

286

Great Basin Naturalist Memoirs

No. 12

Fig. 51. Geographic distribution of Flexamia ^raminea (•), clayi (■), and satilla (D).

inadequate to discriminate between these two
andropogonoid grasses.

Remarks and diagnosis. — We are unaware
of any region in which graininea, clayi, or
satilla occur together. As in the case of
sandersi and delongi, they may represent vi-
cariant populations that have become repro-
ductively isolated and diverged morphologi-
cally. The initial description of clayi from
Kentucky, and subsequent collections of the
species from Georgia and Tennessee, ob-
scured the extent of the range o( clayi. Our
collections of the species from Ohio and the
Maryland and West Virginia Alleghenies, and
identification of material from Maine and New
Hampshire as clayi present a different pic-
ture, portraying the species as a cool-temper-
ate species of the eastern mountains. This
contrasts with that o{ graminea, whose range
is essentially the tall-grass prairie, and satilla,
whose known range is the Gulf Coast (Florida
and south Georgia to Mississippi).

Members of this complex can be readily
separated from other prairiana group species
by aedeagal characters. The aedeagal shaft of
clayi is much more expanded basad of the
gonopore than the usual condition in grami-

nea. However, some specimens oi graminea
from south Texas have a basally expanded
gonopore that could be confused with that of
clayi. The most consistent character that sepa-
rates clayi and graniinea is the condition of the
anteapical lobes of the apodemal processes. In
clayi the processes each have a prominent
anteapical lobe, which is directed mesad. The
shaft in satilla is much narrower than that of
clayi.

44. Flexamia satilla Hamilton & Ross

Flexamia satilla Hamilton & Ross 1975: 606.

Important CHARACTERS. — Length of c? 3.4
(3.3-3.4) mm, of 9 3.6 mm. Crown produced
(1.49 X interocular width; 0.74 x head width).
Face variable, but without distinct interocular
band; usually dark brown or nearly entirely
black. [6] Genitalia much as in clayi, but with
aedeagal shaft ( Fig. 44J) not as expanded at
base of gonopore. [9] Sternum VII as for
group; ovipositor with base of first valvula
rounded, with recurved portion broad and
extending dorsad.

Geographical distribution. — The type
locality in south Georgia is just north of

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Whitcomb, Hicks: Flexamia Revision

287

Okefenokee Swamp. We have found other
specimens, collected by Beamer, from the
Gulf Coast in Florida and Mississippi.

Biology, — This species was collected on an
andropogonoid grass, probably Andropogon
virginicus (K. G. A. Hamilton, personal com-
munication).

Remarks and diagnosis. — This species is
sympatric with producta in the Gulf Coast
region, and two specimens had been filed in
the KU collection in the producta drawer.
These specimens were tentatively separated
from producta by the face, which was black,
without a conspicuous interocular band.
Unfortunately, some specimens with equally
dark faces turned out, when they were
cleared, to be producta. It is therefore easy to
understand how this species escaped recogni-
tion for many years.

The degree of genitalic difference between
clayi and satilla falls well within that consid-
ered by some workers to represent geo-
graphic variation. However, we feel that
satilla is subject to the same phenological con-
straints as producta. The hypothesis that the
graminea complex in the East is divided into
Appalachian and Gulf Coast species seems
reasonable, and, given our ability to distin-
guish the species on (admittedly minor) mor-
phological grounds, justifies retention of
satilla.

This species is safely separated from its
sympatric cohort, producta, only by genitalic
examination, although the latter species (un-
like satilla) often has a distinct interocular
band. For distinctions within the clayi com-
plex, see the comparable section under clayi.

Biology of Flexamia

Flexainia species are residents of North
American grasslands, where they specialize
predominantly on dominant, warm-season
grasses (Panicoideae and, especially, Chlori-
doideae). Several species, however, appear to
breed on sedges (Cyperaceae) or rushes (Jun-
caceae). The genus is divided into 13 species
groups; host choice is related to membership
in these groups.

No Flexamia species are known to be uni-
voltine; as a result, they are inhabitants of
mesic grasslands and are poorly adapted for
semiarid grasslands. The habitats of the genus

include true prairie, short-grass plains,
semitropical grasslands, high desert plains or
montane grasslands, riparian grasslands, and
saline or nonsaline lakeshores.

In semiarid grasslands, grass hosts that sup-
port Flexainia species tend to be capable of
growth with minimal precipitation, but are
unlikely to undergo repeated episodes of dor-
mancy through the summer. Perhaps this is
why, despite their proclivity for chloridoid
grasses, not a single Flexamia species special-
izes on Hilaria or Sporoholus species. This is
true despite the fact that some members of
these grass genera are among the major domi-
nants in southwestern grasslands and com-
prise a significant fraction of the regional
grassland biomass. [Note: As a control to this
"natural experiment," Hilaria and Sporoholus
species have acquired specific non-Flexamia
cicadellid guilds of their own (Whitcomb et al.
1987).]

The relation of Flexamia species to their
hosts ranges from habitat specialization (in
primitive species), to various degrees of inter-
and intrageneric oligophagy, to absolute mon-
ophagy. Narrow oligophagy is the most com-
mon strategy. Body size of Flexamia species
appears to be correlated with size of the host
plant. Intraspecific variation in body size may
reflect host plant quality; individuals from
populations at the range periphery are often
small.

Flexamia species respond to several aspects
of grassland structure. Disturbances such as
mowing or burning tend to reduce popula-
tions or to extirpate them altogether; for this
reason Flexamia is an excellent genus for as-
sessing habitat quality of grassland reserves
(Whitcomb 1987). On a local level, popula-
tions of Flexamia species are almost always
much higher in more mesic segregates of
grasslands defined by minor topographic vari-
ation than in adjacent xeric sites (Whitcomb et
al. 1987). On a regional level the degree of
fidelity to single hosts appears to be least in
mesic prairie or, perhaps, in southeastern
grasslands (Whitcomb et al. 1987). The struc-
ture of grassland dominance hierarchies may
determine host choice.

Habitat patchiness has had a profound ef-
fect on Flexamia species on both historical and
contemporary time scales. For monophagous
species, their hosts are evolutionary islands.
Contemporary host patchiness may affect

288

Great Basin Naturalist Memoirs

No. 12

presence or absence of Flexamia species
within their ranges. However, no Flexamia
species is able to track its host throughout the
entire range of the host. This not only is the
result of patchy host occurrence at the range
periphery, but is often reinforced by climate.
In many instances host colonization appears to
be blocked by climatic factors. For example,
the Flexamia complex that colonizes side-oats
grama {Bouteloua curiipendula) is divided
into six species, none of which is known to
overlap in range. The isolating mechanisms
are those associated with the different climatic
regimes of prairie, Chihuahuan and Sonoran
desert, interdesert grasslands of the Gila
Mountains of southeastern New Mexico, and
the eastern and western Mexican grasslands.
Buchloe dactyloides , Schizachijrium scopar-
ium, and Miihlenbergia porteri are similarly
partioned, as is Bouteloua gracilis (although
less spectacularly so). The possible role of host
biotype (in its inherent genetic composition),
apart from the purely phenologically deter-
mined availability of growing host, has not
been determined.

We have seen no evidence that Flexamia
species are highly fecund; they are rarely cap-
tured at lights or in aerial traps. There is no
evidence of seasonal movement under condi-
tions that permit observation of immigration
of Athysanella macropters or Laevicephalus
adults into suitable grassland. We therefore
predict that the life history strategy of
Flexamia species will be found to resemble
K- rather than r-selection (MacArthur and
Wilson 1967).

Eggs of most species are presumably
oviposited in host tissue. In temperate re-
gions it is reasonable to assume that eggs laid
in the field in autumn are in diapause. In
central latitudes first-generation adults ap-
pear in early June. In subtropical latitudes,
such as south Florida and south Texas, collec-
tion records from December, January, and
February indicate that reproduction may oc-
cur throughout the year.

Species Concept in Flexamia

The species concept that emerges from our
treatment of Flexamia is a composite of indi-
vidual decisions concerning species. We faced
diflPicult decisions in several Flexamia species
groups.

In the prairiana group there are three
sets of close sisters. In one of these sets
{prairiana -reflexa), the types differ so sub-
stantially that no immediate problem is evi-
dent. When specimens from different geo-
graphic regions were examined, however,
problems emerged. The holotype of prairi-
ana, from Illinois, represents the northeast-
ern periphery of the range of this species (Fig.
46). Females from the southwestern part of
the range, especially those from the montane
grasslands of the Chihuahuan Desert region
or the Trans-Pecos shrub savanna, differ from
northern individuals in certain features of the
first valvulae; these populations colonize
Bothriochloa spp. rather than little bluestem
{Schizachyrium scoparium). However, we
found no characters of the male genitalia that
distinguish the Bothriochloa populations.
Further, as discussed by Young and Beirne
(1958), the male aedeagal characteristics of
reflcxa and prairiana vary intraspecifically. It
appears that both species, although wide-
spread, are distributed patchily in some parts
of their ranges. Under these circumstances it
is reasonable to suspect the existence of repro-
ductively isolated populations. For example,
we would not be surprised if the Chihuahuan
populations of prairiana proved to be repro-
ductively isolated from the tall-grass prairie
populations.

Our decision to retain species concepts pro-
posed by Ross and Cooley (1969) and Hamil-
ton and Ross (1975) defining the sandersi-de-
longi and satilla-clayi sister pairs has been
discussed under the species descriptions of
delongi and satilla. In each case we elected to
retain the newly described species on the
criterion of probable phenological isolation
and consistency of (admittedly) minor mor-
phological differences.

The Anasazi form of arenicola presents a
more difficult problem. This population, cen-
tered in the Four Corners area of the South-
west, is reproductively isolated from popula-
tions oi arenicola in the Nebraska Sand Hills.
Further, it has a distinctive morphological
characteristic — the unpaired, aedeagal pro-
cess is invariably broken, presumably in copu-
lation. Although this character tends to define
the Anasazi form, we ha\e found one speci-
men oi arenicola from Nebraska with a broken
aedeagus. Thus, despite our strong suspicion

1988

Whitcomb, HickS; Flexamia Revision

289

that a certain amount of morphological diver-
gence has occurred between the two popula-
tions, we have found no consistent character
to separate the two populations. Because we
feel that a taxonomic designation is inappro-
priate, we refer to the Four Corners popula-
tion simply as the Anasazi form o{ arenicola .

The celata-stylata and beaincri-texana sis-
ter pairs also merit brief mention. The distinc-
tive process of the male pygofer that occurs in
both members of the celata-stylata set sug-
gests that the divergence is relatively recent.
Because the area of present-day occurrence of
celata was largely created by the most recent
glacial maximum (Wright 1970), we at first
thought that this speciation event was Holo-
cene in origin, as the minor degree of morpho-
logic divergence would suggest. The geo-
graphic proximity of the ranges of the two
species seemed to suggest that speciation oc-
curred by dispersal of stylata or a similar an-
cestor into sandhill grasslands that emerged
postglaciation. However, the morphological
evidence contradicts this tidy explanation.
The aedeagal symmetry o( celata and the pres-
ence of presumably plesiomorphic, tapered,
notched plates similar to those of arenicola or
decora demand that celata be considered the
most closely related to the ancestral lineage.
Perhaps during various glacial episodes sandy
grasslands were much more common than
they are today. The existence of three other
uncommon sand-inhabiting Flexamia species
{grammica, arenicola, and areolata), in com-
bination with other studies (Whitcomb et al.
1986, 1987, 1988) indicating that only domi-
nant grasses accumulate specialists, supports
this hypothesis.

The divergence of texana and beameri is
also unclear. This murkiness is perhaps an
artifact of the rarity of the species; each is
known from single collections of E. D. Ball
and R. H. Beamer, respectively. We there-
fore know little of the ecology of these species.
Each of these species occurs just outside the
periphery of the known range of inflata, in
one case in a cold region in New York, in the
other a hot region of south central Texas. The
propensity of inflata to form ephemeral, iso-
lated colonies on a variety of hosts (Whitcomb
et al. 1986) ought, theoretically, to spawn pe-
ripheral new species. Although beameri and
texana appear to be closely related, the bio-
geographic data suggest that an independent

origin for the two species from the inflata
lineage should not be completely discounted.

Species problems in the prairiana and flex-
iilosa groups, though interesting and chal-
lenging, pale when compared to the pectinata
group. This group, prior to this study, con-
sisted of only two species. We now report
expansion of the group to nine species. Three
of the new species (mescalero, jacala, and
collorum) differ profoundly from previously
recognized species. The remaining four spe-
cies {gila, bandarita, minima, and zamora)
are closely related members of a complex that
includes the previously recognized pectinata
and doeringae. It is likely that all of the spe-
cies are specialists of side-oats grama {Bou-
teloiia curtipendula). This grass host is impor-
tant (in many regions dominant) throughout
most of the prairie, from North Dakota to
Ohio to Kansas to north Texas to eastern New
Mexico; throughout this region of importance
the characters of the styles, aedeagal tips, and
female sternum VII of pectinata are relatively
constant. Similarly, populations of doeringae
in southeastern Arizona (where the climate
has a strong Sonoran influence) show an in-
traspecifically constant profile in the morphol-
ogy of the aedeagus, style, and female ster-
num VII. We now report discovery of a
species (bandarita) from the Chisos Moun-
tains and Marathon Basin of Texas which, in
some respects, is intermediate between pecti-
nata and doeringae. Flexamia bandarita oc-
curs in an area that is rich in endemism, as
attested by our description herein ofzacate, a
new Chihuahuan sister of the Sonoran can-
yonensis.

We were delighted to find this endemic
until we examined Flexamia specimens taken
from side-oats grama in the intervening re-
gion between the Chisos Mountains and
southeastern Arizona that could be referred
to neither bandarita nor doeringae. Rather,
they possessed a mosaic of characteristics rep-
resentative not only of bandarita and doe-
ringae, but of pectinata as well. With collec-
tions from three widely separated localities in
New Mexico and Chihuahua, Mexico, totaling
eleven individuals, we were compelled to rec-
ognize another new species (gila).

Although we are uncomfortable separating
a large complex occurring on a single host
into a number of species, we would be even
more uncomfortable illustrating the variation

290

Great Basin Naturalist Memoirs

No. 12

within a single hypothetical polymorphic
"species" and attempting to explain why we
included such divergent characters under a
single name. In our view such a treatment
would require redefinition of the species con-
cept in deltocephaline leafhoppers.

Once we recognized the existence of a spe-
cies complex of side-oats grama specialists, it
was easier for us to accept the results of our
reexamination of mini7na DeLong & Hersh-
berger and zatnora DeLong & Hershberger
from Mexico. In these examinations we were
assisted by the availability of eight previously
unidentified specimens referable to minima
from Monterrey, Mexico, a location consider-
ably north of the type locality. These speci-
mens and the type of minima reveal a mor-
phologic profile similar in some respects to
zamora and pectinata, but with substantial
differences in the male plates and styles and
the female sternum VII. Similarly, on the ba-
sis of the female sternum VII and, to a lesser
extent, the male plates and styles, zatnora can
be separated from all other members of the
complex.

Construction of a map (Fig. 15) of the geo-
graphical distribution of members of the com-
plex clarifies the taxonomic status of the spe-
cies. The distribution of the complex, like that
of the host (side-oats grama), encompasses a
large percentage of North American grassland
area. However, the six species of the complex
occur in very different vegetational regions,
some of which are recognizable at the biome
level. Furthermore, side-oats grama is rare in
the Texas mesquite-acacia savanna, so there is
an essential discontinuity in host distribution
that probably inhibits gene flow between the
United States and Mexican populations of
side-oats grama specialists. Part of the barrier
between pectinata and minima and zamora is,
in fact, the area occupied by doeringae, han-
darita, and gila. Therefore, the existence of
reproductively isolated populations of Flex-
amia specialists on side-oats grama in grass-
lands of the United States and Mexico is prob-
able, and the reinstatement of minima and
zamora is justified.

The concept of climatically limited special-
ists that geographically partition a single grass
host is supported to some extent by studies in
Mexico (Triplehorn and Nault 1985) of Dal-
bulus species, most of which appear to be
Tripsacum specialists. Further support is

given by the recent demonstration (Hicks et
al. 1988) that in New Mexico blue grama
(Bouteloua gracilis) is colonized by seven
Athijsanella species, but that the distribution
of each species is confined to regions that
are readily interpretable in terms of recog-
nized biogeographic regions (e.g., Colorado
Plateau, high plains, short-grass prairie,
Chihuahuan grasslands). We believe that
Bouteloua curtipendula is also partitioned by
its Flexamia specialists.

In an ideal world we would have postponed
all decisions concerning this complex and in-
stead organized an expedition to the South-
west and Mexico to collect Flexamia from
side-oats grama. However, publication of the
conclusions herein need not discourage such
exploration; perhaps it will encourage it.

In summary, we have defined Flexamia
species as populations with consistent and
readily demonstrable morphological differ-
ences whose reproductive isolation from pos-
sibly conspecific populations can be inferred
on biogeographic grounds.

Phylogeny of Flexamia

Young and Beirne (1958) established the
groundwork for a Flexamia phylogeny by their
careful descriptions of genitalic and other
characters of the genus. Also, although they
did not propose a formal phylogeny, they dis-
cussed species relationships at some length.
With a minimum of assumptions, these pro-
posals and relationships can be presented as a
cladogram (Fig. 52).

H. H. Ross was intrigued by Flexamia and
in 1970 published a preliminary phylogeny of
the genus in which the species were identified
only by number and no apomorphies were
described. Although Ross did not present
an explicit proposal, his unpublished notes,
which specify apomorphies and synapomor-
phies, make it possible to diagram his phy-
logeny also (Fig. 53).

OuTGROUPS — Ross considered Spartopyge
Young & Beirne and Aflexia Oman to repre-
sent Flexamia outgroups. Young and Beirne
(1958) felt that Aflexia was not especially
closely related to Flexamia. Although the
habitus of Alapus DeLong and Sleesman is
very different (Beamer and Tuthill 1934) from
that o{ Spartopyge (Fig. 54), its genitalic char-
acters (Figs. 55F,G) suggest relationship with
this genus.

1988

Whitcomb, Hicks. Flexamia Revision

291

•I s

•S ■§

■§ i

\ y

ANCESTOR

Fig. 52. Species relationships in Flexamia : proposal of Young and Beirne (1958). These authors proposed an ancestral
condition in which the Flexamia aedeagus possessed two pairs of recurved, bilaterally symmetrical appendages at or
near the apex of the aedeagal shaft. From this ancestral condition, the authors proposed three or four independent lines
of development, without specifying a phylogenetic sequence in which the lines diverged. In one line of development
(I), the apical pair of processes tended to fuse and the anteapical to remain separate {picta and pyrops). In a second line
of development (II), the pairs of processes are (1) either preserved, as in surcula and curvata , or (2) the apical processes
are reduced and the basal processes absent (abbreviata), or both processes are absent (3) but the location of the
gonopore (in camjonensis) suggests a derivation from the condition in abbreviata. Young and Beirne suggested that
pectinata and doeringae, lacking any trace of aedeagal processes, represent the culmination of one of the developmen-
tal lines. We take the liberty of placing these two species (4) at the culmination of line II. In the final lineage the
important event was (5) the development of an unpaired ventral aedeagal process. In IIIA this occurred either (6) with
additional processes, as in serrata , or (7) with branching of more apical processes, as in slossonae and albida . In albida
(8) the unpaired ventral process is adherent to the shaft. The final sublineage (IIIB) encompasses most species of the
genus. In the more generalized species, the gonopore is small, not elongated, and located on the caudoventral surface
of the aedeagus near the apex (9). Young and Beirne placed seven species {decora, shjlata, modica, ritana, arizonensis,
flexulosa, and inflata) in this cluster. In prairiana a phyletic torsion of the shaft has occurred (10) so that the gonopore
is anteapical on the anterodorsal surface of the shaft. This species has a pronounced groove on the unpaired process,
which in imputans constitutes the gonopore (11). In reflexa the unpaired process is fused basally to the aedeagal shaft
(12) but is free at its extremity; the gonopore is circular, but at some distance from the base of the process. In sandersi
the gonopore has become elongate (13). In producta the movement of the gonopore, to the left side of the dorsal surface
of the shaft, far removed from the apex, approaches a maximum (14). In graminea and clayi the unpaired process has
almost completely disappeared and the gonopore is spiral (15). In grammica, dakota, and atlantica the anteapical
processes have been lost; these species were tentatively placed in line IIIB.

292

Great Basin Naturalist Memoirs

No. 12

o g 5

2 S =

E E C -S o.

:2 I a E

Fig. .53. Flexatnia phylogeny proposed by Ross (1970, and unpublished notes). The tree diagram was transposed to a
cladogram. Identification of species and apomorphies used by Ross to construct the cladogram are taken from
unpubhshed notes. The apomorphies and conserved plesiomorphies designated were (1) mesal brush on pygofer; (2)
thin aedeagal shaft with basal processes; (3) lack of aedeagal processes; (4) width of aedeagus, thick (4) or thin (4'); (5)
length of male style; (6) angulate pygofer; (7) four aedeagal processes; (8) aedeagus with mesal process; (9) acute aedeagal
apex; (10) long, mesal aedeagal process; (11) unpaired aedeagal process bifurcate; (12) aedeagal processes twisted; (13)
dorsal location of gonopore; (14) fusion of connective and aedeagus; basal location of median process and gonopore
(somewhat = 15, more so = 16); widening (17) of median process to bean-shaped structure appressed to shaft; (18) loss
of medial process.

Future phylogenetic hypotheses concern-
ing Flexamia should be broadened to include
a much wider range of outgroups than those
discussed above. Candidates for study should
perhaps include Enantiocephalus Haupt, a
Eurasian genus (Figs. 55A,B). K. G. A.
Hamilton (personal communication) has sug-
gested that Mocuellus Ribaut and related gen-
era should not be overlooked as outgroups.
Finally, we will soon add a new genus (in
preparation), presumably also related to Flex-
amia. This genus consists, so far as known, of a
single species that specializes on Muhlenber-
gia arenacea in saline flats in semiarid regions
of New Mexico and Arizona. The former
genus Acurhinus {-^ Donjdium) may be more
distant from Flexamia than once thought (Lin-
navuori 1959). Acurhinus macukitus (Osborn),

known only from three females in the Ohio
State University collection, is currently re-
garded as a species of Hododoecus Jacoli, an
essentially African genus of the Stirellini (Lin-
navuori and DeLong 1978). It is evident that
a phylogeny of deltocephaline genera, espe-
cially those with linear connectives articu-
lated with the aedeagus, would greatly en-
hance all phylogenies of deltocephaline
species.

In this study we have chosen two Sparto-
pijge species [miranda (Knull) and mexicana
(DeLong & Hershberger)] and Alapus elon-
gattis Beamer & Tuthill for outgroup compari-
son.

Phylogeny of Flexamia croups. — We
have developed an intuitive phylogeny utiliz-
ing many of the characters and concepts

1988

Whitcomb, Hicks; Flexamia Revision

293

Fig. 54. Spartopyge Young & Beirne, the principal Flexainia outgroup. Spartopyge mexicana (DeLong & Hersh-
berger): A, habitus; B, face; C, male plates, ventral aspect; D, female sternum VII; E, S. miranda (KnuU), female
sternum VII.

developed by Young and Beirne (1958) and
Ross (1970, unpublished notes). Details of our
reconstruction follow:

A. Designation of sister species. We first
identified synapomorphies that define close

sister species sets. We also assigned polarities
within these sets by defining significant aut-
apomorphies of one of the set members.

The sets and defining synapomorphies [s]
and autapomorphies [au] were:

294

Great Basin Naturalist Memoirs

No. 12

Fig. 55. Male genitalia of genera related to Flexamia. Enantiocephalus corntitus (Herrich-Schaffer): A, aedeagus and
connective, lateral aspect; B, male pygofer, lateral aspect. Spartopyge mexicana (DeLong & Hershberger): C,
aedeagus, lateral aspect; D, male pygofer, lateral aspect. Spartopyge miranda (Knull); E, male pygofer, lateral aspect.
Alapus elongatus Beamer ix Tuthill: F, aedeagus and connective, lateral aspect; G, male pygofer, lateral aspect.

surctila —> curvata [s] Aedeagus with two pairs of pro-
cesses, one apical, one anteapical
(Figs.7F,G).

[au] Apical processes long and not
in bilaterally symmetrical plane
(Fig. 7F). Gonopore an elongate
channel.

host =
Buchloe dactyloides

zacate —* canyonensis [s] Aedeagus curved, acute api-
cally, without aedeagal processes
host - (Figs. 7I,J).

Muhlenbergia porteri [au] Gonopore at midlength of
aedeagal shaft.

ininiiTui —* zamora fs] Style apices in lateral aspect
bluntly produced (Figs. 13A,B).
[au] Outer teeth of female sternum
VII incurved, longer than inner
teeth (Fig. 14B).

host
unknown

bandarita —* gila

host =

Bouteloua

curtipendtila

albida —* slossunae

habitat
specialists

picta —* pyrops

host =
Aristida spp.

[s] Female sternum VII with two
produced, median teeth (Figs.

14D,E). Male plates broad, blunt-
ly rounded (Figs. 11D,E).
[au] Male plates nearly as long as
pygofer (Fig. HE).

[s] Posterior lobe of male pygofer
undifferentiated (Figs. 62Q,T).
[au] Male plates exceed pygofers
(Fig. 8B): female sternum VII
trilobed (Fig. lOB). Aedeagus
and connective partially fused.

[s] Aedeagal tip capitate, toothed,
with pair of subapical processes
(Figs. 7K,L); pygofers (Figs. 5A,
B) angulate with ventral process.
[au] Vertex extensively produced,
anteapical aedeagal processes
(Fig. 7L) more remote from apex,
male plates (Fig. 9B) very short,
rounded.

1988

Whitcomb, Hicks; Flexamia Revision

295

arizonensis —*
Jlexulosa
hosts = Aristidal
Bouteloua g,racilis

youngi —* decora

host =
Muhlerihergia
richardsonis

celata —* stylata

hosts =
Redfieldia/
Muhlenhergia spp.

heameri —> texana

hosts
unknown

[s] Male plates fFi«s. 33A,Bl
short, divergent apically.
[an] Apical aedeagal processes
short (Fig. 3 IK).

[s] Unpaired ventral aedeagal pro-
cesses (Figs. 31F,G) symmetrical
and much longer than paired pro-
cesses.

[an] Unpaired process (Fig. 31G)
without median groove, aedeagal
shaft curved.

[s] Face (Fig. 3B) with discrete
lines forming interocular band;
three aedeagal processes angu-
larly divergent from shaft (Figs.
31H,I). Pygofer with distinct pos-
terior process (Figs. 6200,PP).
[au] Apices of male plates sub-
rectangular; aedeagus (Fig. 31H)
asymmetrical.

[s] Aedeagal processes about
equal in length, but ventral pro-
cesses not in sagittal plane (Figs.
31 A, D).

[au] Aedeagal processes twisted
apically (Fig. 31A).

imputans — » areolata [s] Unpaired process lateral, with

hosts ^

Muhlenhergia

cuspidatal

Eragrostis

spectahilis

ventral groove (Figs. 31B,C).
[au] Aedeagus and connective
fused, unique habitus (Fig. 2F).

})rairiana — » rejlexa
hosts
andropogonoid
grasses

sandersi — » dehmgi

hosts
andropogonoid
grasses

satilla —* claiji
hosts
andropogonoid
grasses

[s] Gonopore dorsiJ (Figs. 44D,G).
[au] Groove absent on unpaired
process (Fig. 44D).

[s] Lateral process appressed to
aedeagal shaft (Figs. 44B,C).
[au] unpaired process (Fig. 44C)
at less acute angle with aedeagal
shaft,

[s] P'lared, elongate gonopore

(Figs. 44H,J).

[au] Gonopore broad basally (Fig.

44H).

B. Designation ofclades. The next step
consisted of appending other species to the
sister sets to define clades:

mtntnia —*

zamora —* pectinata

host probably =

Bouteloua

curtipendula

handarita —* gila —*

doeringae

host =

Bouteloua

curtipendida

graminea -> satilla —
clayi

hosts =

andropogonoid

grasses

[s] Style with apical lobe not con-
stricted (Figs. IIA-C);
[au] Male plates (Fig. IIC) sub-
rectangular; style tip (Fig. 13C)
not produced in lateral aspect.

[s] Middle teeth (Figs. 14D-F) of
female sternum VII longer than
outer teeth.

[au] Middle teeth long, associated
to form process (Fig. 14F).

[s] Spiral gonopore (Figs. 44F,H,J)
[au] Flared gonopore (Fig. 44H)

C. Establishment of species groups. Clades
were next assembled into species groups.
These are the groups (I-XIII) described
herein. Synapomorphies defining clades and
species groups are given in the descriptions to
the groups and in the legends to Figures
56-59.

D. Establishment of polarity between Spar-
topy ge and Flexamia. The dorsal and facial
habitus of grammica (Fig. 2C), albida (Fig.
2A), and Spartopyge (Fig. 54) are clearly ho-
mologous. On the other hand, few characters
of grammica are clearly homologous to those
of other Flexamia species. The morphology of
the male aedeagus and connective places this
species in the genus, but the reduced number
of aedeagal apices and their apparent phyletic
twisting make it difficult to assign homology.
The male pygofer and female sternum VII
suggest retention of plesiomorphic characters
of an ancestor allied to Spartopyge . We there-
fore hypothesize the polarity Spartopyge — ►

grammica . Derivation of the similarly striped
albida group, of course, could have occurred
only after development of paired aedeagal
processes and fusion of the anteapical pair into
a single, unpaired process (Fig. 56). Revers-
ing the polarity between grammica (or albida )
and Spartopyge would require generation of a
wide array of apomorphies present in Spar-
topyge, including a brush of setae on the inte-
rior of the pygofer and the complex structures
of the male genitalia (Fig. 55C), accompanied
by complete loss of all apical processes. Also,
the unique sternum VII (Figs. 54D,E) of
S. mexicana and S. miranda would have to
arise suddenly. Finally, if Spartopyge were
derived from the Flexamia lineage, it would
be necessary to postulate a significant apo-
morphic increase in size. This would consti-
tute a reversal of the overall trend for size
reduction and simplification in the Delto-
cephalinae in general (Whitcomb et al. 1986)
and Flexamia in particular. We therefore

296

Great Basin Naturalist Memoirs

No. 12

=2 5

II III

7 8

VII

Fig. 56. See facing page.

VIII

XII

XIII

21 22

hypothesize (with Young and Beirne, and
Ross) that Spartopyge is an outgroup of the
Flexamia hneage. We beheve, with these
workers, that a primitive Flexamia ancestor
arose from the Spartopyge hneage and later
gave rise to a hneage with paired, aedeagal
processes. This hneage in some way gave rise
to grammica but, ultimately, to all Flexamia
groups IV-XIII. We assume that the dorsal
location of the gonopore in grammica and
prairiana is a result of independent evolution-
ary events, since these species share few other
significant characters. Whether the minute,
paired processes that occur midlength on the
Spartopyge aedeagus are homologous with
the Flexamia aedeagal processes is conjec-
tural.

Phylogeny of Flexamia groups and spe-
cies. The postulated relationships and polari-

ties permit a cladogram to be drawn relating
Flexamia groups and key species to the out-
groups Alapus and Spartopyge (Fig. 56). Sev-
eral significant possible variations of the pro-
posed tree are given in Figure 57. Finally, the
phylogeny of Flexamia species was inferred on
the basis of the characters discussed above.
We present cladograms for groups I-IX (Fig.
58), groups VII-XI (Fig. 59), and groups
XI-XIII (Fig. 60). Synapomorphies used to
define clades within each group are given in
the legends to the figures.

Character transformations. — Our intu-
itive phylogeny provided general guidelines
concerning character transformation in Flex-
amia. From these concepts we constructed
branched transformations for the PHYSYS
program of Mickevich and Farris (1984).
These transformations, which are described

1988 Whitcomb, Hicks Flexamia Revision 297

Fig. 56. Proposed phylogeny of Flexamia groiip.s and key species. Synapoinorphies [s] or, for monobasic groups,
autapomorphies [an] are indicated. We hypothesize an ancestral form (1) that possessed dorsal stripes (Fig. 54 A) and a
pale face with a contrasting black interoeiilar band (Fig. 54B) similar to those of Figs. 3A and 3B, relatively wide male
plates (Fig. 54C); an aedeagus distinctly jointed with the connective, without apical processes, and with an anteapical
gonopore on the caudoventral surface. Divergence of this lineage led (3) to Alapus (Figs. 55F,(v), represented in this
diagram by elongatus, and to Spartopy^e, represented (4) in this diagram by miranda and mexicana, in which [s] the
dorsal aedeagal apodeme is large and hoodlike (Fig. 55C). The transformations between Flexamia and its outgroups are
tenuous (see te.xt). In Flexamia [s] the dorsal aedeagal apodeme (Fig. 6) has a pair of conspicuous appendages directed
caudoventrad or caudad (5). In the first lineage (6) to diverge, the plesiomorphic aedeagal apex, which lacked processes,
was retained, as were the caudoventral position of the gonopore and broad, ancestral plates; the contrasting face was
retained in a minority of species, but the striped habitus was completely lost. Synapoinorphies defining this lineage
include [s] the curved aedeagus lacking processes (Figs. 71, J; 16A, 17A, 2()A) and the male pygofers, which possess both
dorsal and ventral constrictions (Figs. 62A-I). This lineage diverged into the pectinata (7) and the abhreviata-zacate
lineages (8). In the pectinata group the plates (Fig. 11) are distinctively parallel-sided basally (except in collorum). In
the abhreviata-zacate lineage the gonopore was at first elongate (Fig. 24) and [s] the posterior lobe of the male pygofer
strongly produced (Figs. 62H,I). In the ahbreviata group (II) the plates (Fig. 8E) are reduced (9), and [au] a distinctive
pair of minute apical processes (Fig. 7H) developed. The small aedeagal processes of the abbreviata group are not
clearly homologous with other aedeagal processes in Flexamia. In the zacate group (III) the gonopore (Fig. 24) was at
first elongate (10) but in canyonensis moved to the aedeagal midpoint. At this point a significant apomorphy appeared.
An ancestral form (11) is hypothesized in which two pairs of aedeagal processes were present. It is unclear whether this
form had fully developed before divergence of grammica (IV), in which the plesiomorphic feature of dorsal stripes was
retained (12), but in which [au] the aedeagal processes were distinctively modified (Fig. 7E) and the gonopore had
moved to the dorsal surface (12). A possible apomorphy linking g,rammica with the curvata group is the foot-shaped
style apex (see illustrations of Young and Beirne 1958). In the cttrvata-picta lineage [s] both pairs of aedeagal processes
are present (14), but many habitus features, including stripes and contrasting face, have been lost. In the curvata group
(V) two pairs of processes are present (15) and [s] unmodified (Figs. 7F,G). In the picta group (VI) the apical processes
are [s] modified (Figs. 7K,L) as flangelike, toothed appendages (16). Remaining species are defined by an apomorphy
that developed with the presumed fusion (17) of the anteapical processes into a single, unpaired process that was
plesiomorphically located ventrally in the sagittal plane (Figs. 31, 61A). A second apomorphy characterizing this
divergence is the bifurcation (Figs. 7A-C) of the apical aedeagal processes, a feature that was subsequently lost (22).
The plesiomorphic state of dorsal stripes and contrasting face was preserved (18) during the fusion of subapical
processes and is present (Figs. 2A,B) in the albida group (VII). This lineage, which can be defined [s] in terms of the
absence of dorsal or ventral constrictions in the posterior lobe of the male pygofer (Figs. 62Q,T), is represented today by
the distantly related sister species albida (19) and slossonae (20); for autapomorphies defining these species, see text. In
serrata (VIII) [au] additional dorsal aedeagal processes (Fig. 7A) developed (21), and, while the contrasting face was
retained, dorsal stripes were lost (Fig. 1). In ritana (IX) the dorsal stripes were retained (Fig. 2D) but greatly reduced.
This monobasic group (23) is defined [au] by the unique, pointed aedeagal apex (Fig. 7D). In one lineage of the^^xw/osa
group (25), the unpaired aedeagal process is, plesiomorphically, ventral in the sagittal plane, permitting retention of
aedeagal symmetry (Figs. 31G,I,J,K,L). The plesiomorphic pale face with black interocular band (Figs. 3B,F,G) was
also retained. This clade is defined [s] by the triangulate plates (Figs. 33A-F, 34D) with conspicuous, apical divergence.
(But in stylata [Fig. 34E], the plates [au] are truncated.) In the remaining species (26) the unpaired process underwent
phyletic twisting (Figs. 61B-H) and the contrasting pale face, which had persisted through many nodes, was finally lost.
In the inflata subgroup of group X, [s] the displacement of the unpaired process (Fig. 61 B) was relatively small (27), but
[s] in the imputans-areolata (XI-XII) lineage the process had become lateral (Figs. 31B,C) and bore the gonopore as an
elongate slit (29). Before divergence of groups XI-XIII, the connective and aedeagus had remained articulated (but see
discussion of group VI in text). We hypothesize that after divergence of the imputans-areolata lineage, a fusion of
aedeagus and connective occurred (31) in areolata . The imputans-areolata lineage is very specialized in the location of
the unpaired process, in the black face, and the greenish dorsal pigmentation. Of the two species, areolata is the more
highly specialized (Figs. 2F, lOQ); in fact, it may be the most highly specialized of all Flexamia species. It is most
unlikely that the widely distributed and variable prairiana (see text) and the entire prairiana group (XIII) were derived
from the specialist areolata lineage. Instead, we propose that divergence of a second lineage in which aedeagus and
connective fused (32) led to the prairiana group. This fusion, the traces of which are evident (Fig. 6F) in prairiana , is
designated [s] as a synapomorphy defining group XIII, and the fusion in areolata is defined as an autapomorphy
defining that species. Details of species phylogenies are presented in detail in Figures 58-60.

in Appendix II, Part III, involved the follow- ness (lateral aspect) of the aedeagus; (7) length

ing characters: (1) relationship between ae- ofthe aedeagus; (8) form of the aedeagal shaft;

deagus and connective; (2) morphology ofthe (9) form of margins of paired, aedeagal pro-

aedeagal apex; (3) morphology of anteapical cesses; (10) form and position ofthe gonopore;

aedeagal processes; (4) phyletic twisting ofthe (11) morphology of apices ofthe apodemal

unpaired ventral process (Fig. 61); (5) curva- arms; (12) position of apodemal arms; (13)

ture (lateral aspect) ofthe aedeagus; (6) thick- width of dorsal keels of the connective; (14)

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III

VIII

VII

XII

XIII

Fig. 57. Alternative resolutions for Flexamia group phylogenies. A. In this resolution (I) two groups (I and III) are
defined by [s] curved aedeagus without processes (Figs. 71, J, 16A, 2IA). The alternative lineage (2) is defined by an
apomorphic elongation of the gonopore. This was followed by derivation of a further lineage (3) in which aedeagal
processes are present. At first these are [s] minute (II abbreviata. Fig. 7H) but later become (4) fully developed (see
Figure 56). This resolution avoids hypothesizing independent origins for the aedeagal processes in abbreviata and
curvata but hypothesizes additional nodes between Spartopyge and the Flexamia species with very similar habitus,
implying repeated, independent losses of the complex pattern of striping in Spartopyge, grainmica, and albida. B. In
this resolution, it is hypothesized (1) that the serrata group and decora subgroup of group X belong to a common
lineage, and that the inflata subgroup of group X is more closely related to ritana. This construction has considerable
merit and takes into account the similarities between the serrata male plates (Fig. 8C) and those of the decora subgroup
(Figs. 33, 34D, E), but deemphasizes many similarities between members of the inflata and decora lineages, and would
augur for elevation of these lineages to groups. C. It is possible to hypothesize a single (2) fusion of aedeagus and
connective, rather than the two fusions hypothesized in Figure 56. This construction, in our view, gives insufficient
weight to several synapomorphies that link areolata and imputans, which would, in fact, have been designated as a
close sister species set had they not differed in the crucial relationship of aedeagus to connective. See discussion in
legend to Figure 56, and text.

morphology of the male pygofer (Fig. 62); (15)
morphology of the male plates; (16) morphol-
ogy of the female sternum VII; (17) presence
or absence of dorsal stripes; (18) coloration of
the face.

The above transformations, as presented in
Appendix II, are suitable for entry into the
PHYSYS program and should serve as a useful
starting point for more refined analysis of
Flexamia phylogeny.

Summary of phylogenetic model. — In
summary, after numerous iterations, we pro-
pose a formal phylogeny for Flexamia. The
model is rooted in earlier informal work of
Young and Beirne (1958) and Ross (1970, and
unpublished notes). Our model, in addition to
its formality, considers 14 species not avail-
able to earlier workers and proposes species
groupings based in large part on synapomor-
phies. In particular, the addition of seven spe-

cies to the pectinata group (I) and four species
to the flexulosa group (X) adds new phyloge-
netic perspectives.

We have accepted the hypothesis of Young
and Beirne of an ancestral form with two pairs
of aedeagal processes. We also endorse the
proposal of Young and Beirne (and Ross) that
the anteapical pair of processes fused to form a
single, unpaired process (Fig. 52). Further,
we accept the proposal of Ross (Fig. 53) that
the fusion of aedeagus and connective is an
important synapomorphy that defines the
prairiami group.

The informal proposal of Ross differed from
that of Young and Beirne in one major way (in
addition to its presentation as a tree). Ross
believed that the lack of aedeagal processes in
pectimita and canyonensis was plesiomorphic
and related it to the similar lack of processes in
his outgroups, Spartopyge and Aflexia. In our

1988

Whitcomb, Hicks: Flexamia Revision

299

proposal we have sided with Ross. Unless dif-
ferent outgroups were chosen, we feel such an
assignment is inevitable. Also, transforma-
tions within Flexamia are best explained by
the chosen polarity. For example, viewing the
pectinata group as the terminus of a simplifi-
cation of the aedeagus would require the
transformation of male plates from narrow to
broad, in direct contrast to the general ten-
dency for streamlining of the plates that runs
throughout the remainder of the Flexamia
tree.

We have assumed, with Ross, that the
minute processes in abbreviata are not ho-
mologous with other Flexaryiia aedeagal pro-
cesses. This assumption, which permits many
other reasonable character transformations to
be postulated, appeared in our model only
after numerous iterations failed to find a bet-
ter resolution, considering all transforma-
tions. Such issues are best addressed by close
examination of the structures themselves,
preferably by scanning electron microscopy,
for evidence that would bear on the question
of homology. It is possible that a computer-
ized search for a parsimonious tree would turn
up a cladogram that avoids the pitfalls we
encountered in our intuitive search. In any
event, we present in Figure 57A an alterna-
tive cladogram that, whatever its other prob-
lems, assumes homology between the aedea-
gal processes of abbreviata, grammica, and
curvata.

Young and Beirne (1958), although uncer-
tain about the status oidakota, atlantica, and
grajnmica, nevertheless placed them tenta-
tively in their lineage IIIB (Fig. 52). We con-
cur with Ross in his placement of dakota and
atlantica. However, we disagree with both
Young and Beirne and Ross in the placement
oi grammica. Whereas the dorsal location of
the gonopore and the presence of two serrate,
aedeagal processes of this species is clearly
highly apomorphic, we see nothing else in
grammica that links it to group XIII, in which
the dorsal location of the gonopore is a signifi-
cant apomorphy. For example, the male
plates and pygofers, the female sternum VII,
the clear articulation of aedeagus and connec-
tive, the striped habitus and pale face, and the
overall large size oi grammica are derivable as
transformations from the outgroup Spar-
topyge, but they would produce chaos if in-
serted into transformations in group XIII (or

any other group or groups between VII and
XII). In fact, the ties between Spartopyge and
grammica are sufficiently reasonable that we
have couched our argument for the polarity
Spartopyge — ► Flexaitiia in terms oi grammica
(see earlier section).

The model we present solves many of the
problems that we encountered. Resolution of
many (juestions was made possible by propos-
ing (1) that aedeagal processes arose in the
course of Flexamia evolution and were not
inherited from an unknown ancestor; (2) that
aedeagal symmetry was plesiomorphic (but
note the apomorphic symmetry in dakota).
Further character transformations have been
proposed (Appendix II, Figs. 61, 62) that lead
to consistent hypotheses for transformation of
other characters (e.g. , bases of the first valvu-
lae, which are presented in Fig. 63 in terms of
our proposal but were not used to derive it).
The proposed resolution avoids hypothesizing
independent acquisitions of the unique
striped habitus present in Spartopyge, and in
grammica, albida, and slossonae. It further
avoids hypothesizing independent deriva-
tions of the unique facial habitus of the out-
group, which was retained through a consid-
erable segment oi Flexamia evolution. We do
hypothesize several independent losses of
these distinctive sets of habitus characters.
Finally, the proposed model follows the fol-
lowing general polarities: (1) tropical grass-
lands — ► subtropical grasslands — ► temperate
grasslands — ► [semiarid grasslands, prairie];
(2) large — ► small size; (3) grass generalist — ►
chloridoid specialist — ► mixed chloridoid-pan-
icoid specialist — > andropogonoid specialist;
(4) striped habitus —* [light habitus (semiarid),
brown (prairie)]; and (5) aedeagal symmetry
and caudoventral gonopore — ► highly special-
ized aedeagal configurations and gonopore lo-
cations.

But not all the problems have been solved.
We regard the cladogram for groups I-III to
be tentative. The position of mescalero as a
member of group I is unclear; this unique
species -might have been more appropriately
considered to be a separate group linking
groups I and III. In fact, the general question
of polarity of the pectinata group should be
critically addressed. We are not certain how to
interpret evidence of considerable recent spe-
ciation in this group (i.e., bandarital gila , min-
imal zamoral pectinata). It is likely, given our

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•S

■§

•£

.§

••2

■*

Cfc

■O

ea.

9 10

2 I

III

24 25

2<i

L'9

Fig. .58. See facing page.

discovery of many new species from the
Southwest, that further acquisitions of Flex-
amia or related genera from Mexico may
change our perceptions dramatically. Also, a
wider selection of outgroups would clarify the
status of the early Flexamia lineages.

Young and Beirne (1958) proposed that
grooves oil the unpaired aedeagal process
were vestiges of a plesiomorphic condition in
which the halves of the processes were sepa-
rate. This condition occurs in several species
{youngi, imputans, prairiana), but we have
been unable to link these species in any trans-
formation that is consistent with other trans-
formations.

Finally, we propose, with some confidence,
that the articulation between aedeagus and
connective was subject to fusion or partial

fusion in four nodes of our model. These were
in the lineages leading to picta-pyrops, to are-
olata, to slossonae, and to the prairiana
group. Given the different products of each of
these fusions, and the position on the tree in
which they occurred, we have no problem in
treating each as a synapomorphy.

The genus Flexamia is rich in characters
and should provide an excellent model for the
PHYSYS program. It is likely that careful
study and numerous iterations using such a
program will further improve phylogenetic
hypotheses concerning Flexamia.

blogeographic implications of the
Phylogenetic Model

Ross (1965, 1970) felt that many speciation

1988 Whitcomb.HicksFlexamia Revision 301

Fig. 58. Phylogeny of species of groups MX. The following events, inelnding synaponiorphies [s] and aiitaponior-
phies [au] are postulated; (1) In the plesiomorphic state of the pectinata-abhrcviata subtree, the male plates were broad
(Fig. 11), the male pygofers constricted (Fig. 62F), the gonopore oval on the caudoventral surface, and the aedeagus
(Figs. 12A-H) without apical processes; (2) in one subtree of the pectinata group (comprising eight of its nine species),
the aedeagal tip remained unmodified but flared (Figs. 12A-H). In all three species of the gi/a division (3) of this group,
the female sternum VII developed [s] long, median, paired teeth (Figs, 14D-F); in doeririf^ae (4) these teeth [au]
became associated into a medial process (Fig. 14F). In gila (5) [au] the male plates are elongate (Fig. 1 1 F), in handtirita
(6) [au] shortened (Fig. II D). Evolution of this cluster, oi' pectinata, and, perhapsalsoof muiima and zamora. occurred
on Bouteloua curtipendnla. The three species of the minima cluster are defined [auj by (8) subrectangular plates
{pectinata: Fig. IIC), (9) short plates {minima: Fig. IIA), and (10) outer teeth of hind margin of female sternum VII
longer than median teeth {zamora: Fig. 14B). Divergence from this cluster (1 1) involved [s] development of avicephali-
form style tips. In coUorum (12), which appears to be a specialist of Bouteloua uniflora, a close relative of B.
curtipendula, [au] the plates (Fig. IIG) are extremely short. In jacala the plates (Fig. IIH) remained long and broad
(13); this is the only Flexamia species with both long, broad plates and avicephaliform style tips. In another species
cluster a tendency toward development of aedeagal processes developed. In mescalero (15) [au] the aedeagus (Fig. 22)
is slightly capitate rather than flared. This specialist oi Muhlenhergia pauciflora retained relatively broad plates (Fig.
Ill) and constricted pygofers (Fig. 62B) typical of the pectinata group; we therefore regard it as a monobasic subgroup
of group I. A monophyletic cluster of three species is presumed to have arisen (16) from this lineage. Each of these
species has [s] a pygofer with a conspicuously produced posterior lobe (Figs. 62H,I). In abbreviata (group II) the
aedeagal apex [au] bears (Fig. 7H) two minute, paired processes (17). In this species and in zacate (18) the gonopore
(Fig. 24) is elongate. F. zacate is the only species of the genus that lacks aedeagal processes but has an elongate
gonopore. In canyonensis the gonopore [au] is located (19) approximately at the midpoint of the aedeagal shaft,
presumably as a consolidation of the elongate gonopore type of zacate. The shape [au] of the aedeagal shaft in
camjonensis (Fig. 6C) is also unique. The sister species zacate and canyonensis are both specialists of Muhlenber^ia
porteri. In a very important divergence (20), development of two pairs of aedeagal processes is hypothesized. In
grammica (22), a Calamovilfa specialist, the plesiomorphic habitus of the Spartopyge dorsum and face were retained.
Two of the four hypothesized aedeagal processes may have been lost. The pygofers and male plates also suggest
relationships with Spartopyge , but in contrast to this outgroup, aedeagal processes are present. The relationship of
grammica to other Flexamia species with aedeagal processes is unclear, given the [au] specific asymmetry of these
processes (Fig. 7E) and the location of the gonopore, which has moved to a dorsal position. It is possibly significant [s?]
that the style apices of the grammica and curvata groups are foot-shaped (21). In the curvata set (23) of sister species
{Buchloe specialists) [s] two pairs of processes are present. In curvata (25) [au] the gonopore is elongate. In the picta
lineage {Aristida specialists) two pairs of processes are present but [s] the apical pair has been modified into flangelike,
toothed structures (26). Several autapomorphies [au] characterize (28) pyrops (see text). From this subtree the
remaining species of Flexamia were derived; this derivation was featured (29) by an apomorphic fusion of the anteapical
pair of processes into a single, unpaired process located on the ventral surface of the aedeagus in the sagittal plane.

events involving phytophagous insects could The progression of evolutionary divergence
be explained by host transfer; the genus Flex- in Flexamia can best be described in terms of
amia was one of the genera that he chose for the phylogeny we propose. In Figures 64-67
the study of host choice and speciation in the we illustrate the postulated biogeographic
Cicadellidae. Ironically, Ross happened to and host transfer events,
work largely in central, northern, and south- We believe that the most closely related
eastern grasslands. In these regions the study outgroup is Spartopijge. We have no informa-
of cicadellid host relationships is more diffi- tion on the host relationships of 7ne.ricana, but
cult than in the Southwest, and, as it turned miranda occurs on Bouteloua curtipendula in
out, almost all of the undescribed species in Arizona. The general facies o{ Spartopyge ap-
the genus occurred outside the geographic pears to be homologous with that of some
regions in which he conducted his extensive Flexamia groups (albida, grammica, and ri-
fieldwork. tana groups). We propose that diversification
Today, in retrospect, we report that the of the Spartopyge lineage in Mexican grass-
choice of Flexamia was a wise one; however, lands led to Alapus and to Flexamia.
fewer speciations appear to be due to host We propose that the initial divergence (Fig.
transfer than Ross might have predicted. Sis- 65) of the Flexamia lineage resulted from spe-
ter species have in many cases proved to be cializationofageneralist ancestor on side-oats
specialists of the same host or closely related grama, Bouteloua curtipendula. This lineage,
hosts. Indeed, entire groups of species have influenced by host patchiness and/or regional
proved to have similar (often congeneric) pat- climatic differences, diverged by dispersal or
terns of host selection. vicariance into at least six species. These are

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VII

VIII

14
12

10 11

9 20

XII

XIII

Fig. 59. Phylogeny of species of groups VII-XI. The following events, including synapomorphies [s] and autapomor-
phies [au] are hypothesized. Plesiomorphically, this cluster retained (1) certain features of the Sparlopyge habitus (i.e.,
dorsal stripes and a pale face with black interocular band). However, prior to divergence, two pairs of aedeagal
processes had developed, and, subsequently, the anteapical pair had fused to form a single, unpaired ventral process
(Figs. 7A-C, 31). The lineage diverged into the (2) albida (VU), (5) serrata (VIII), and (6) ritana (IX) groups. The
relationships of these groups is discussed in the legend to Figure 56. A large clade that then diverged (8) was associated
mainly with Muhlenbergia. In all species of this subgroup the plesiomorphic character of a pale face with black
interocular band was retained, but the dorsal stripes were lost. The subgroup is defined [s] by the triangulate male
plates (Figs. 33A-F, 34D,E), which are strongly divergent apically (but see serrata [Fig. 8C]; the plates of this species
suggest a link to the decora subgroup). In the ijoung,i sister species set (host: Muhlenbergia richardsonis ), the unpaired
ventral process (Figs. 3IF,G) retained its plesiomorphic position and length (20), but in the remaining species [s] the
process was shortened (9). In modica the aedeagus is subsymmetrical (11) with [au] the short, unpaired process (Fig.
31M) slightly displaced from the sagittal plane and associated with the shaft basally; the pygofer is relatively
undifferentiated. In the lineage that diverged subsequently, the pygofer (Fig. 62) became substantially modified (10).
We hypothesize that it first became ovate, a condition that still persists [au] in arenicola (16). In stylata and celata [s] it
developed [s] a distinctive posterior process (17: see te.xt for further synapomorphies), and in arizonensis (au) it
developed a conspicuous ventral process (14). The male plates oiflexidosa and arizotwnsis (Figs. 33A, B) are in each case
(13, 14) di.stinctive [au]. In the inflata subgroup of group X (24), the beginning of phyletic torsion of the aedeagus is
evident (Fig. 61 B). In this subgroup the face is modified; traces of the interocular band are present in some individuals,
but, when present, it is brown and contrasts much less sharply with the remainder of the face. The plates are short, and
[s] the dorsal margin of the pygofer is essentially flat, giving the pygofers a boatlike outline in lateral aspect (Figs.
62GC,HH,II). Two apparent (but see text) sisters, texana and beameri, are identified by [au] unique aedeagal
configurations (Figs. 31A,D). From the moderately displaced aedeagus of the inflata group, further torsion (29) led to
groups XI-XIII.

1988

Whitcomb, Hicks: Flexamia Revision

303

Si
I

o
■4i

>4J

-§

■§

XII

18 19

XIII

Fig. 60. Phylogeny of species of groups XI-XIII. Synapomorphies [s] and autapomorphies [au] are identified. In the
plesiomorphic state (1) there may have been moderate aedeagal asymmetry, as in inflata (Fig. 61B). In the imputans-
areolata hneage (2) [s] the unpaired processes are lateral, bearing a slitlike gonopore (Figs. 31B,C). In imputans (3) the
ancestral articulation between aedeagus and connective was retained. This unique species is defined by a suite of
apomorphies, including its black face, greenish dorsal pigmentation, and lack of dorsal markings, but lacks an
autapomorphy. In areolata (Fig. 61C) [au] the aedeagus and connective have fused, and a conspicuous, black spot (Fig.
2F) is present in the corium (4). Because the imputans and areolata lineage is defined by several synapomorphies (see
text), we have designated them as sister species, despite the important difference in the state of articulation between
the aedeagus and connective. We hypothesize a second, independent fusion of aedeagus and connective that serves as
a synapomorphy [s] defining group XIII (4). In the prairiana-reflexa lineage [s] the gonopore has become dorsal (Figs.
61D,E), as has the unpaired process (6). In prairiana (7) the traces of this fusion [au] are present as a transverse line
(Fig. 6F) on the surface of the aedeagal shaft. The aedeagal configuration [au] is also unique (Fig. 61D). In reflexa (8) the
gonopore [au] has become associated with the unpaired process (Fig. 61E). In atlantica one process has been lost, and
[au] the aedeagus (Fig. 44A) has become highly asymmetrical (9). In producta [au] the gonopore has moved (10) to a
lateral position (Fig, 61G), whereas in dakota (11) the gonopore remained dorsal (Fig. 441), but the unpaired process
has been lost. The symmetry of the dakota aedeagus is therefore apomorphic. In the sandersi-delongi lineage (12) the
unpaired process has been retained but is present [s] as a lateral process that is closely appressed to the aedeagal shaft
(Fig. 61F). In all three species of the final lineage (15), the unpaired process has essentially disappeared; instead, [s] the
gonopore has become spiral, with [au] a narrow, basal portion (Fig. 44F) as in graminea (16), or [s] with a wider base (17)
as in clayi and satilla (Figs. 44H, J). For discussion of the minor morphological divergences that define the sister species
of the sandersi-delongi and clayi-satilla sets, see text.

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Fig. 61. Postulated character transformation: phyletic twisting and modification of the Flexamia aedeagus. In decora
(A) a plesiomorphic state exists in which the unpaired process is ventral and in the sagittal plane; the aedeagus is
therefore symmetrical. In inflata (B) there is a small displacement of the unpaired process. In areolata (C) the unpaired
process is lateral and bears the gonopore. In prairiana (D) the process is dorsal, as is the gonopore. In reflexa (E) the
gonopore has become associated with the dorsal unpaired process. In sandersi (F) the unpaired process is reduced and
appressed to the shaft. In producta (G) the process is reduced or absent and the gonopore is lateral. In ^raininea (H) the
process has disappeared entirely, the apical processes are reduced, and the gonopore is spiral. Aspects; AD,
anterodorsal; CV; caudoventral; DS, dorsal; LT, lateral; VN, ventral. Aedeagal structures redrawn from Young and
Beirne(1958).

adapted to northeastern {minima) and west
central (zamora) Mexican grasslands, to Chi-
huahuan (bandarita ) and Sonoran {doeringae)
montane grasslands of the Southwest, to the
interdesert montane region of the Gila Moun-
tains igila), and to the tall-grass and mixed-
grass prairie of the United States (pectinata).
Also derived from the side-oats lineage were
jacala (Mexico: host unknown) and collorum
(Edwards plateau; host: Bouteloua uniflora),
a close relative of side-oats grama). Clearly,
most or all of the nodes in the pectinata lin-
eage are Mexican or southwestern; none rep-
resent events that assuredly took place north
of the Mexican-U.S. border.

A second lineage of the pectinata subtree
resulted in the divergence of mescalero, a
distinctive species that represents a host
transfer to New Mexican muhly, Midilenher-
gia pauciflora. Another branch of this lineage
transferred to temperate Bouteloua grass-
lands of Mexico. One of the branches (abbre-
viata group) inhabited the mixed Bouteloua
grasslands of Mexico and, by gradual adapta-
tion, to similar grasslands of the United
States. Another branch adapted to semiarid
grasslands by colonizing bush muhlenbergia
{Muhlenbergia porteri)., divergence of this
lineage led to the contemporary canijonensis
(Sonoran) and zacate (Chihuahuan).

Consider next the surcula-curvata lineage.
Both species possess the four aedeagal pro-
cesses that Young and Beirne proposed as ple-
siomorphic; both species are Buchloe special-

ists. We propose that an ancient Mexican an-
cestral line adapted to B. dactyloides, which
led to the present-day surcula and, later, to
curvata. This divergence (Fig. 64) may have
been mediated by climate. The grasslands of
northeastern Mexico, where surcula may
have arisen, have escaped major disturbance
since the Cretaceous (Rosen 1978). This re-
gion has an extremely warm, temperate or
subtropical climate and a Chihuahuan precip-
itation pattern, both of which differ im-
mensely from the dry, temperate grasslands
where curvata occurs today.

Another branch of the curvata subtree di-
verged, perhaps in Mexico or south Texas,
leading to the picta-pyrops sister set. These
species are Aristida specialists.

One of the major events in Flexamia phy-
logeny was marked by the fusion of the an-
teapical pair of aedeagal processes into a sin-
gle, mesal process. The most primitive
species in which this feature is retained are
members of the albida, serrata, and ritana
groups. Young and Beirne argued (and Ross
concurred) that slossonae, albida, and serrata
were related. We agree but consider their
divergence to be an ancient one, given their
entirely different climatic reciuirements (sub-
tropical, prairie, and northern grasslands, re-
spectively), and their retention of the Spar-
topyge habitus type. Perhaps the divergence
of the albida lineage into four species, each
with imiciue climatic adaptations, may have
occupied a distinct stage in Flexamia evolu-
tion.

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Whitcomb, Hicks: Flexamia Revision

305

Fig. 62. Character transformation of the Flexamia pygofer. Diagrammed Hneage was based in part on phylogeny of
groups (see Figure 56). A, collorum; B, mescalero; C, handarita; DJacala; E, gila; F, pectinata, G, doeringae; H,
ahhreviata- I, canyonensis; J, pijrops- K, curvata, L, surcula- M, picta; N. Spartopijge miranda; O, grammica; P,
serrata,- Q, a/foida; R, A/apus elongatus; S, Spartopyge iijexicana; T, .s/ossoriae; U, ritana; V, impt^fans; W, prainana;
X re/Zexa- Y af/anfica, Z, areo/ata, AA, producta; BB, ffa/coto; CC, sanrfersi; DD, c/ayi, EE, graminea, ¥¥ decora;
GG, inflata; HH, texaria, II, foeani^ri, JJ, modica, KK, yowngj, LL, arenicola; MM. ari^onenm; NN,/^cxu/osa; OO,
sfy/ato; PP, celata . Some pygofers were redrawn from Young and Beirne (1958) and Lowry and Blocker (1987).

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Fig. 63. Bases of first valvulae of Flexamia species. The depicted tree represents phylogeny presented in Figures
56-60. A, doeringae; B, gila; C, handarita; D, colloruni; E, mescalero; F, curvata; G, pectinata; H, canyonensis; I,
surcuhi; J, pictoi K, zacate; L, Alapus elongatiis; M, abbreviata; N, Spartopyge mexicana- O, albida, P, serrata; Q,
decora; R, modica; S, arenicola; T, inflata; U, slossonae; V, ritana; W, ijoungu X, celata; Y, arizonensis; Z, beameri;
AA, stylata; hB, flexulosa; CC, texana ,DD, atlantica; EE, producta; FF, sandersi; GG, c/ai/i, HH, reflexa; II,
imputans; JJ, prairiana, KK, dakota; LL, graminea.

Interestingly, there are present-day clues
to the feeding strategies of this primitive com-
plex. We suspect that slossonae and alhida are
habitat specialists, whereas serrata special-
izes on Muhlenbergia richardsonis . Ancestral
Flexamia species in Mexican grasslands may
have been general grass feeders; this strategy
has been partially retained by some lineages,
including those that retained the plesiomor-
phic striped habitus.

Perhaps the striped habitus is optimal for
habitat specialists. Loss of the habitus oc-
curred homoplastically several times; in at
least one case (serrata), it accompanied host
specialization in northern, temperate grass-
lands (on Muhlenbergia richardsonis). This
event could actually have taken place far south
of the current range oi serrata during a glacial

maximum in the early Pleistocene. Muhlen-
bergia richardsonis not only occurs as far
north as Wood Buffalo National Park in
Canada, but also occurs in Mexico, pre-
sumably as a relict. Throughout much of the
northern part of its range, this warm-season
grass is an "island" surrounded by a sea of
cool-season grasses. Such a condition is ideal
for evolution of a specialist lineage from a
more general ancestor; taxonomic distinctive-
ness has been recognized as an important
force in determining insect species richness
on plant hosts (Lawton and Schroder 1977). In
any event, morphological evidence indicates
that a subtree o( Flexamia evolved on Muhlen-
bergia. The first event (Fig. 66) appears to
have been the divergence of the serrata
and decora lineages. The decora subgroup is

1988

Whitcomb, Hicks: Flexamia Revision

307

Fig. 64. Geographic interpretation of Flexamia phylogeny. Early evolution is postulated to be Mexican. The
Spartopyge lineage diverged into mexicana and miranda. From this lineage Flexamia diverged. The first major node
involved the divergence of the pectinata group, perhaps as a result of specialization on Boutelotia curtipendula. This
divergence is hypothesized to have occurred in the grasslands of northeastern Mexico, a region thought to have
remained stable over recent geologic history (Rosen 1978). Subsequent divergences, which occurred in the same
general region of Mexico, were those of the abbreviata-zacate and curvata-picta lineages. The abbreviata lineage
adapted to Boutelona spp., the curvata group to Buchloe dactyloides, and the picta group to Aristida spp. The
divergence of the albida lineage, characterized by a single, plesiomorphically ventral aedeagal process, may have
occurred somewhat north of the Sierra Madre Orientale region, perhaps in Texan grasslands. Many conteinporary
species emerged from this lineage. The most plesiomorphic of these (members of the albida group) are relatively
uncommon today; albida and slossonae do not appear to be strict host specialists, a feeding habit that we hypothesize to
be plesiomorphic for the genus. Actual divergences in temperate zones may have occurred farther south than
diagrammed, if they occurred during periods of glacial advance.

presumed to have divided by vicariance dur-
ing glacial episodes into a Rocky Mountain-
Canadian {decora) lineage, and a Great Basin
lineage (yoiingi), both specialists on M.
richardsonis . Alternatively, a second lineage
developed, leading to a New Mexican mon-
tane lineage (modica) that transferred from
Muhlenhergia richardsonis to Muhlenbergia
repens. Further divergence led to arenicola
(host: Muhlenbergia pungens) and to a set of

sister species (celata-stylata) that specialized
on Muhlenhergia or Redfieldia (and possibly
other glasses of sandy grasslands), respec-
tively. Finally, this line appears to have trans-
ferred from Muhlenbergia to the vast resource
of Bouteloua gracilis in the plains region of
New Mexico. This resource has been shared
with abbreviata, which arrived in the New
Mexican plains by an entirely different geo-
graphic route (i.e., Mexican mixed Bouteloua

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Fig. 65. Geographic interpretation of phylogeny of the pectmata group. A focal origin in grasslands of eastern Mexico
is hypothesized, A western lineage comprised gila, handarita, and doeringae. This lineage speciated exclusively on
Bouteloua curtipendtda. Morphological data suggest that mescalcro, which apparently transferred to Muhlenbergia
pauciflora, belongs on a deep branch. The subtree consisting of za»iora, minima, and pectinata is closely related to the
doeringae subtree; like all members of that subtree, pectinata specializes on B. ctitiipendiila (side-oats grama). F.
coUorum may have been derived from a lineage on B. curiipendula, since its host is B. uniflora, which is closely related
to side-oats grama. The host ofjacala is unknown.

grasslands). (Interestingly, in contrast to
Athysanella species [Hicks et al. 1988], both
Flexamia blue grama specialists colonize this
host throughout most of its range in New Mex-
ico). A sister species, arizonensis, transferred
to another dominant southwestern grass,
Aristida purpurea. We emphasize that con-
siderable climatic diversity exists today in the
Southwest and no doubt has existed through-
out the Pleistocene and Holocene as the result
of elevational gradients (Axelrod and Raven
1985, Cole 1986, Cronquist 1978, Thompson
and Mead 1982, VanDevender and Spaulding
1979). Therefore, evolutionary processes that
might take place only over wide geographic

distances in regions of relatively constant ele-
vation may be focused in small geographic
areas in the Southwest.

Another lineage, possibly more closely al-
lied to the ritana than serrata lineage, pre-
sumably diverged in grasslands of the south-
ern plains. This is the inflata subgroup of the
flexulosa group. One contemporary represen-
tative (inflata) exhibits one of the widest host
ranges in the genus; largest populations occur
on Muhlcnhcrgki aspcrifolia in wetland areas.
If this apparent preference indicates an ances-
tral Muhlenbergia host, it would be necessary
to postulate a monumental subsequent
widening of host range to pooid grasses or

1988

Whitcomb, Hicks: Flexamia Revision

309

Fig. 66. Geographic interpretation of phylogeny of the serrata, ritana, flexulosa, imptitans, and areolata groups.
Texan grasslands may have been a focal area for initial divergence of lineages. F. serrata appears to be a Muhlenbergia
specialist. The ritana group appears to have diverged in the Southwest. The contemporary distributions of the albida,
serrata, and ritana groups suggest that they may have speciated by vicariant mechanisms or, in the case oi serrata, by
a combination of vicariance and host transfer. An eastern lineage consists oiinflata and beameri. From a biogeographic
perspective, it is most reasonable to propose an independent origin oitexana and beameri, each from inflate or a closely
related lineage. The present-day distribution oiinflata suggests that it diverged in cool, temperate grasslands; Texas, of
course, was cool during glacial maxima. In general, events postulated as northern in these reconstructions could have
taken place much farther south during glaciations. The imputans-areolata lineage appears to have diverged in the
prairie, on Muhlenbergia cuspidata ind Eragrostis spectabilis , respectively. The western lineage of the/7ex«/osa group
may be related (see Fig. 59) to the serrata group; if this association can be confirmed, the derivation of the decora set of
sister species from the serrata lineage would make an attractive hypothesis from a biogeographic standpoint, since all of
the species (including youngi) specialize on Muhlenbergia richardsonis. Derivation oi modica (host: Muhlenbergia
ripens) and arenicola (host; Muhlenbergia pungens) represents further evolution that occurred by host transfers within
Muhlenbergia. The derivation of flexulosa and arizonensis by host transfer may have been a consequence of the
regional abundance (in New Mexico and eastern Arizona) of their hosts, Bouteloua gracilis and Aristida purpurea,
respectively; presumably, the region was colonized from the north. Finally, derivation of celata (host: Redfieldia
flexuosa and perhaps other sandhill grasses) and stylata (hosts: Muhlenbergia spp.) is hypothesized to have followed a
high-plains route around the eastern edge of the high Rocky Mountains.

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Fig. 67. Geographic interpretation of the phylogeny of the prairiana group. Although present-day distributions and
host preferences do not offer obvious hypotheses for speciation mechanisms, an origin for the prairiana group in Texan
grasslands is hypothesized. From this region prairiana presumably moved northward in postglacial episodes, where it
adapted to little bluestem (Schizachyrium scoparium ) or other andropogonoid grasses. In the tall-grass prairie, reflexa
diverged from this lineage; this species often seems to be associated with big bluestem (Andropogon gerardii),
particularly in glades of the eastern United States. F. atlantica may have transferred in postglacial times from
chlorodoid grasses, on which it predominates in the Southwest, to switchgrass, Panicum virgattim. This host has been
a conduit for colonization by atlantica of the eastern United States, including the Atlantic coastal prairies, where its host
achieves high importance. Derivation oi sandersi may have occurred by transfer to Andropogon virginicus, the first
colonist in early fire succession in the Southeast. From this lineage, presumably post-Wisconsinan, delongi may have
diverged by colonization of little bluestem in the glaciated area northwest of Illinois. Specialization on Schizachyrium
scopariuin and subsequent divergence led to dakota in the dry prairies of the plains, to graminea in Texas and
northward through the tall-grass prairie, and to a lineage of eastern species. These species, also residents of
andropogonoid grasses, are residents of the Gulf Coast (satilla), or the Appalachian mountains to Maine and cool,
temperate savannas of Illinois and Ohio (clayi).

Juncus. It may be more likely that the cur-
rently observed host range breadth repre-
sents simply a retention of plesiomorphic gen-
eralism. In any event, this lineage has
spawned two additional species: texana (south
central Texas) and beameri (west slope of the
Adirondack Mountains, New York). The cir-
cumstances surrounding these divergences
are unclear, since we have no data on their

host relationships.

After divergence ofinflata, another lineage
diverged in the prairie, leading to imputans
(probable host: Muhlenbergia cuspidata) and
areolata (host: Erag^rostis spectabilis) on one
hand, and to the prairiana group on the other.

All speciation events of the prairiana group
(Fig. 67) occurred either in prairie, savanna,
or southeastern grasslands. We hypothesize

1988

Whitcomb, HickS: Flexamia Revision

311

an ancestor for this group that was adapted to
both chloridoid and panicoid grasses. From
such an ancestor, prairiana (liost: Schiza-
chyrium or Bothriochloa) and reflexa (host:
Andropogon gerardii) were derived, as were
atlantica (hosts: chloridoid grasses and/or
Panicum virgatum), and prodiicta (hosts
chloridoid grasses, or [possibly] a habitat spe-
cialist). Finally, specialization on Schiza-
chyrium scoparium and/or Andropogon vir-
ginicus led (Fig. 67) to lineages that adapted to
the most xeric of Andropogon habitats (da-
kota, a Schizachyriiim specialist of the west-
ern Great Plains), graminea (a Schizachyriiim
specialist of the tall-grass prairie), and the
sa7idersi-delongi and satilla-clayi sister pairs
of eastern andropogonoid grasslands.

Stages of Speciation

Mayr (1942) believed that at any given time
all stages of evolution should be observable.
We believe that this is indeed the case in the
genus Flexamia. We recognize at least 10 lev-
els in the process of speciation in Flexamia:

1. Nonisolated ephemeral populations: (i)
Many populations of Flexamia inflata appear
to be ephemeral. These populations often oc-
cur on pooid grasses such as Poa or Festuca;
presence of immature insects confirms ovipo-
sition; (ii) F. abbreviata colonizes Miihlenber-
gia richardsonis in the Sacramento Moun-
tains of New Mexico and M. reverchonii in the
Edwards Plateau of Texas, although in the
latter case we have not confirmed oviposition.

2. Nonisolated stable populations: (i) F. in-
flata consistently colonizes path rush (Juncus
tenuis) in Illinois and Maryland; populations,
including immatures, are found regularly on
that host. This is an especially interesting
case, since it may represent an interfamily
transfer by a species that has limited
oligophagy. (ii) Flexamia atlantica regularly
colonizes large plantings of weeping love grass
(Eragrostis spectahilis) and bermudagrass
{Cynodon dactylon) in Oklahoma and Texas.
(iii) Similarly, prodiicta colonizes bermuda-
grass and Zoysia japonica in Maryland. These
are essentially "natural experiments " that may
mimic to a large degree, but in a shorter geo-
logic time period, the effect of radical changes
in the dominance hierarchy of grasses induced
by cyclic climatic change.

3. Disjunct populations that have not
diverged morphologically: In the following

cases we have found no morphological varia-
tion associated with disjunct populations, (i) A
disjunct population oi dakota, a specialist on
Schizachyriiim scoparium in the western
Great Plains, occurs on that host in the Loess
Hills of western Iowa, (ii) Flexamia reflexa
occurs (always, it seems, in association with
big bluestem, Andropogon gerardii) in iso-
lated grassland islands in the eastern decidu-
ous forest as far east as the Soldier's Delight
serpentine barrens in Baltimore County,
Maryland.

4. Isolated stable populations with minor
morphological divergence: In each of the fol-
lowing cases the populations may or may not
represent biological species, (i) Two disjunct
populations (Nebraska Sand Hills and Four
Corners or Anasazi populations) of arenicola
have been discovered; the unpaired, aedeagal
processes of the Anasazi population always
break, whereas the processes of the Sand Hills
population rarely break. Other characters
have not been found to separate these popula-
tions, each of which specializes on Muhlen-
bergia pungens. The host population is also
separated into disjunct populations, (ii) The
Mammoth Lakes population oiyoungi, prob-
ably isolated in eastern California in a region
rich in endemics, has diverged somewhat
from the population in the eastern Great
Basin, (iii) Populations of prairiana on Both-
riochloa in the Southwest have begun to di-
verge from the prairie Schizachyriiim popula-
tions, perhaps as the result of phenological
isolation. In this case divergence of female
rather than male structures may signal the
emergence of a new species.

5. Recent (Holocene) speciations: (i) The
separation of delongi from the sandersi lin-
eage may represent a postglacial transfer from
Andropogon virginicus to Schizachyrium sco-
parium in the glaciated region of the northern
prairie, (ii) The divergence ofclayi and satilla
may reflect phenological isolation between
Gulf Coast and Appalachian populations.

6. Pleistocene speciations: Biological and
biogeographic circumstances strongly suggest
that decora and youngi represent vicariant
populations of a common lineage that di-
verged during Pleistocene climatic cycles, re-
taining Miihlenbergia richardsonis.

7. Vicariance and semiisolation: In the case
of Mexican and Southwestern grasslands, spe-
ciation was perhaps achieved by vicariance

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Great Basin Naturalist Memoirs

No. 12

during long periods of semiisolation. It is pos-
sible, in the case of certain species, that there
may have been a punctuated infusion of genes
into semiisolated populations as the climate
became more mesic (Milstead 1960) and the
distance between host stands of side-oats
grama decreased. In any event the emergence
of gila, doeringae, bandarita, zamora, min-
ima, and pectinata, closely related species
that appear to partion the range of side-oats
grama, must have entailed different specia-
tion mechanisms than those described above
and are clearly enforced today by phenologi-
cal isolation.

8. Distinct species with obvious sisters: We
consider the sister pairs of picta-pyrops, ciir-
vata-surcida, canyonensis-zaccite , and prairi-
ana-reflexa to be obvious sisters that occur in
ambiguous circumstances that suggest no par-
ticular age for their divergence. In some cases
(surcula-curvata [tropical/temperate] and za-
cate-canyonensis [Chihuahuan/Sonoran]) the
biogeographic circumstances for the diver-
gences are clear.

9. Species with distant sisters: We consider
the albida-slossonae sister pair to represent a
historically remote divergence. Flexamia slos-
sonae is a subtropical species that has many
autapomorphies separating it from other Flex-
amia species. However, common characters
of habitus and male genitalia leave no doubt
that it is a sister to the tall-grass prairie species
albida.

10. Orphans: The characteristics of gram-
mica are sufficiently unique that it must be
considered an orphan within the genus. In the
absence of a contemporary link, the relation-
ships to other extant species of the genus are
obscure.

From the above examples we feel that our
experience with Flexamia elegantly confirms
Mayr's hypothesis.

Conclusions

The initial choice of Flexamia as a model for
the study of the role of host plants in insect
evolution was a fortuitous one. Whereas some
speciation events in Flexamia must have oc-
curred by host transfer, others surely oc-
curred by vicariance. Further, there is a ten-
dency of individuals representing advanced
lineages to be reduced in size and complexity,
and to be more highly specialized in their host

choice than their ancestors. There is therefore
an immense stake in arriving at a well-sub-
stantiated phylogenetic hypothesis for the
genus that is supported by independent lines
of evidence. In particular, if advances in the
study of molecular evolution prove to be
adaptable to the study of small insects, we
believe that Flexamia would provide an excel-
lent model for study. Because the knowledge
of Flexamia hosts is now adequate, obtaining
live material of most species for such a study
would be an achievable task. The rewards of
further detailed studies would be many but
would include, especially, important insights
into the evolution of life-history strategies of
sap-sucking insects.

When we began our studies of Flexamia,
whose species number had remained rela-
tively constant for a number of years, we did
not dream that there were many remaining
species awaiting discovery. It is difficult for us
to guess, given the results reported here, how
far we have fallen short, even now, of a com-
plete inventory. Our discovery of the role of
climate in inducing and reinforcing cicadellid
speciation makes it clear that the biogeo-
graphically rich areas of the Southwest may
well have many interesting stories that still
await us.

In particular there may well be Mexican
species awaiting discovery. For example, our
phylogenetic hypothesis suggests that there
may have been at one time a species with the
plesiomorphic dorsal stripes of Spartopyge,
grammica, and albida, but which possessed a
full complement of four aedeagal processes.
Because of the compelling logic of all phyloge-
netic schemes for its existence, such a species
seems so real to us that we have jokingly re-
ferred to it as ancestris. Is it possible that
this species, or a closely related descendant of
it, may actually exist in an unexplored Mexi-
can grassland? It is questions such as these
that have driven us to return repeatedly to the
grasslands of the Southwest, which prove to
be every bit as full of enigmas as they are of
answers.

Acknowledgments

There is a Japanese proverb that advises
those who drink water from a well today to
remember the ancestors who dug the well.
The Flexamia well has been dug with many

1988

WniTcoMB, HiCKS; Flexamia Revision

313

spades. The early biological work of E. D.
Ball, H. Osborn, D. M. DeLong, and R. H.
Beamer is implicit in our work. H. H. Ross
introduced us to the genus, framed many of
the questions that we discuss, and left much
unpublished material from his grasslands
(GL) project. D. A. Young, whose 1958 mono-
graph with B. P. Beirne continues to be an
important tool for students of Flexamia,
kindly permitted us to redraw many struc-
tures from drawings in his monograph. We are
particularly pleased to acknowledge the ad-
vice and encouragement of P. W. Oman,
given especially to the senior author during
his early years at Beltsville. Further, a large
fraction of USNM Flexamia had been col-
lected and correctly determined by Oman;
this material was extremely valuable in our
study. Earlier studies on Flexamia at
Beltsville were assisted by M. E. Coan, J. P.
Kramer, and, in later years, J. E. Lowry,
whose illustration oLserrata is presented here
as Figure 1. All other drawings were done by
Debi Roney. Robert Almond and Edward
Clark gave advice and assistance on the graph-
ics. Host records were retrieved from a com-
puterized index developed by D. E. Lynn.
T. Mogel assisted with the scanning electron
microscopy. Assistance with phylogenetic
programs was provided by Mary Mickevich.
We acknowledge further the assistance of
G. W. Byers, P. S. Cwikla and C. P. Triple-
horn, M. W. Nielson, and T. J. Henry and
R. C. Froeschner, who made available the
Flexamia collections of Kansas University,
Ohio State University, Monte Bean Life Sci-
ence Museum (Brigham Young University),
and the United States National Museum, re-
spectively. K. G. A. Hamilton made available
the Flexamia material from the Canadian Na-
tional Collection and provided a wealth of
stimulating comments on the manuscript.
M. SchaufF made many useful comments on
the manuscript. So many friends have helped
in our field studies that all cannot be acknowl-
edged. We are especially indebted, however,
to Kelly Allred of New Mexico State Univer-
sity for continuing lessons on New Mexico
agrostology. Charles T. Mason, Jr., assisted
with grasses of Arizona. R. Chambers of the
U.S. Soil Conservation Service drove us to
the type locality of collortwi and called our
attention to the host. Mike Fleming of
Big Bend National Park and Jim Walters of

Guadalupe National Park assisted field stud-
ies there, and Barton Warnock identified
many plant specimens from Big Bend Na-
tional Park and the Davis Mountains. Riley
Nelson of BYU provided collections from the
Colorado Plateau. P. Cubbage assisted with
fieldwork. Judith Leach, wife of the senior
author, assisted during many field trips and
also made valuable editorial comments on the
manuscript. The editorial staff of The Great
Basin Naturalist helped in many unusual
ways to assure timely completion of the paper.
Finally, H. D. Blocker suggested that the
time for this paper had come, sorted more
than 1,000 unidentified Flexamia from the KU
cicadellid collection, provided Flexamia ma-
terial and records from the KSU collection,
read various drafts of the manuscript, and en-
couraged us in countless other ways. To all
these workers, and many that remain un-
named, we offer our sincerest gratitude.

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